GIFT  OF 


v 
O   <S 


UNIVERSITY    OF    CALIFORNIA    PUBLICATIONS 

IN 

ZOOLOGY 

Vol.  11,  No.  14,  pp.  377-510,  pis.  21-24, 5  text  figs.    February  27,  1914 


A    DETERMINATION   OF  THE   ECONOMIC 
STATUS  OF  THE  WESTERN  MEADOW- 
LARK  (STURNELLA  NEGLEC1A) 
IN  CALIFORNIA 


BY 

HAROLD  CHILD  BRYANT 


A  Thesis  presented  to  the  Faculty  of  the  College  of  Natural 
Sciences  of  the  University  of  California  in  partial  fulfillment 
of  the  requirments  for  the  Degree  of  Doctor  of  Philosophy. 
May,  1913. 


UNIVERSITY  OF  CALIFORNIA  PRESS 
BERKELEY 


UNIVEESITT  OP  CALIFORNIA  PUBLICATIONS 

Note. — The  University  of  California  Publications  are  offered  in  exchange  for  the  publi- 
cations of  learned  societies  and  institutions,  universities  and  libraries.  Complete  lists  of 
all  the  publications  of  the  University  will  be  sent  upon  request.  For  sample  copies,  lists 
of  publications  or  other  information,  address  the  Manager  of  the  University  Press,  Berkeley, 
California,  U.  S.  A.  All  matter  sent  in  exchange  should  be  addressed  to  The  Exchange 
Department,  University  Library,  Berkeley,  California,  U.  S.  A. 

OTTO  HARRASSOWITZ,  B.  FBIEDLAENDEB  &  SOHN, 

LEIPZIG.  BEBLIN. 

Agent  for  the  series  in  American  Arch-  Agent  for  the  series  in  American  Arch- 
aeology and  Ethnology,  Classical  Philology,  aeology  and  Ethnology,  Botany,  Geology, 
Education,  Modern  Philology,  Philosophy,  Geography,  Mathematics,  Pathology,  Physi- 
Psychology,  History.  ology,  Zoology,  and  Memoirs. 

ZOOLOGY.— W.  E.  Bitter  and  C.  A.  Kofoid,  Editors.  Price  per  volume,  $3.50.  Commenc- 
ing with  Volume  n,  this  series  contains  Contributions  from  the  Laboratory  of  the 
Marine  Biological  Association  of  San  Diego. 

Cited  as  Univ.  Calif.  Publ.  Zool. 
Volume  1,  1902-1905,  317  pages,  with  28  plates  $3.50 

Volume  2  (Contributions  from  the  Laboratory  of  the  Marine  Biological  Associa- 
tion of  San  Diego),  1904-1906,  xvii  -f  382  pages,  with  19  plates $3.50 

Volume  S,  1906-1907,  383  pages,  with  23  plates  $3.50 

Volume  4,  1907-1908,  400  pages,  with  24  plates  „.  $3.50 

Volume  5,  1908-1910,  440  pages,  with  34  plates  $3.50 

Vol.  6.  1.  (XXHI)  On  the  Weight  of  Developing  Eggs.  Part  I,  The  Possible 
Significance  of  Such  Investigations,  by  William  E.  Bitter;  Part  II, 
Practicability  of  the  Determinations,  by  Samuel  E.  Bailey.  Pp.  1-10. 
October,  1908  10 

2.  (XXIV)  The  Leptomedusae  of  the  San  Diego  Begion,  by  Harry  Beal 

Torrey.    Pp.  11-31,  with  11  text-figures.    February,  1909  20 

3.  (XXV)  The  Ophiurans  of  the  San  Diego  Begion,  by  J.  F.  McClen- 

don.    Pp.  33-64,  plates  1-6.    July,  1909  SO 

4.  (XXVI)  Halocynthia  johnsoni  n.  sp.:  A  comprehensive  inquiry  as  to 

the  extent  of  law  and  order  that  prevails  in  a  single  animal  species, 

by  Wm.  E.  Bitter.    Pp.  65-114,  plates  7-14.    November,  1909  .50 

5.  (XXVII)  Three  Species  of  Cerianthus  from  Southern  California,  by 

H.  B.  Torrey  and  F.  L.  Kleeberger.     Pp.   115-125,   4  text-figures. 
December,  1909 - 10 

6.  The  Life  History   of   Trypanosoma  dimorphon  Dutton   &   Todd,   by 

Edward  Hindle.    Pp.  127-144,  plates  15-17,  1  text-figure.    December, 
1909 50 

7.  (XXVIH)  A  Quantitative  Study  of  the  Development  of  the  Salpa 

Chain  in  Salpa  fitsiformis-runcinata,  by  Myrtle  Elizabeth  Johnson. 

Pp.  145-176.     March,  1910  35 

8.  A  Bevision  of  the  Genus  Ceratocorys,  Based  on  Skeletal  Morphology, 

by  Charles  Atwood  Kofoid.    Pp.  177-187.    May,  1910 10 

9.  (XXIX)  Preliminary  Report  on  the  Hydrographic  Work  Carried  on  by 

the  Marine  Biological  Station  of  San  Diego,  by  George  F.  McEwen. 

Pp.  189-204;  text-figure  and  map.    May,  1910  15 

10.  (XXX)  Biological  Studies  on  Ccrynorpha.    in,  Regeneration  of  Hy- 

dranth  and  Holdfast,  >y  Harry  Eeal  Torrey.    Pp.  205-221;  16  text- 
figures. 

11.  (XXXI)  "^o1e  on  Geotropism  in  Corywiorpha,  by  Harry  Beal  Torrey. 

Pp.  223-224;  1  text-figure. 

Nos  10  and  11  in  one  cover.  August,  1910 20 

12.  The  Cyclostomatous  Bryozoa  of  the  West  Coast  of  North  America,  by 

Alice  Robertson.    Pp.  225-284;  plates  18-25.    December,  1910  60 

13.  Significance  of  White  Markings  in  Birds  of  the  Order  Passeriformes, 

by  Henry  Chester  Tracy.    Pp.  285-312.    December,  1910  25 

14.  (XXXIII)  Third  Report  on  the  Copepoda  of  the  San  Diego  Region,  by 

Calvin  Olin  Esterly.    Pp.  313-352;  plates  26-32.    February,  1911 40 

15.  The  Genus  Gyrocotyle,  and  Its  Significance  for  Problems  of  Cestode 

Structure  and  Phylogeny,  by  Edna  Earl  Watson.    Pp.  353-468;  plates 

33-48.    June,  1911 - 1-°° 

Index,  pp.  469-478. 

*  Roman  numbers  indicate  sequence  of  the  Contributions  from  the  Laboratory  of  the 
Marine  Biological  Association  of  San  Diego. 


UNIVERSITY    OF    CALIFORNIA    PUBLICATIONS 

IN 

ZOOLOGY 

Vol.  11,  No.  14,  pp.  377-5 10,  pis.  2 1-24,  5  text  figs.    February  27,  1914 


A    DETERMINATION    OF    THE    ECONOMIC 
STATUS  OF  THE  WESTERN  MEADOW- 
LARK  (8 TURN ELL A  NEGLECT  A) 
IN  CALIFORNIA 

BY 

HAROLD  CHILD  BRYANT 


CONTENTS 

PAGE 

Preface   378 

Introduction    382 

History  of  methods  in  economic  ornithology  389 

Investigation   of    the    economic    status    of    the   western   meadowlark 

in  California  395 

A  comparison  of  methods  in  economic  ornithology  397 

The  western  meadowlark  (Sturnella  neglecta)  400 

Field   investigation    401 

Abundance    402 

Nesting   habits   404 

Depredations    404 

Experiments  on  captive  birds  409 

Amount  of  food  required  410 

Time  of  digestion  412 

Examination  of  the  stomach  contents  413 

Collection  and  preservation  of  material  413 

Material    415 

Examination  of  stomach  contents  416 

Identification  of  stomach  contents  420 

.  Food  of  the  western  meadowlark  in  California  420 

Vegetable  food  420 

Animal    food    426 

Inorganic   matter  436 

Principal  articles  of  diet  437 


378          University  of  California  Publications  in  Zoology    [VOL.  11 

PAGE 

Examination  of  feces 437 

Quantity  of  food 440 

Capacity  for  good  or  evil  as  evidenced  by  the  number  of  birds 

taking  different  kinds  of  food 444 

Pood  of  nestlings  447 

Variation  of  kind  of  food  448 

Variation  in  food  according  to  time  of  year  450 

Variation  in  food  habits  according  to  locality 454 

Influence  of  age  and  sex  on  quantity  of  food  455 

Combination  of  field  and  laboratory  work 456 

The  relation  of  birds  to  insect  outbreaks  456 

Verdict  of  ranchers  as  to  the  value  of  the  western  meadowlark 462 

A  determination  of  the  economic  status  of  the  western  meadow- 
lark  in   California   466 

Suggestions  for  the  protection  of  crops  474 

Recommendations  as  to  legislation  475 

Some  interesting  side-lights  on  the  investigation  477 

Parasitism    477 

Malformation    477 

Albinism     478 

Incubation  and  moult  478 

The  effect  of  systematic  destruction  on  the  numbers  of  meadow- 
larks    478 

Natural  death-rate  479 

Do  protective  adaptations  of  insects  protect   them  from  the   at- 
tack of  birds?  480 

Availability  as  a  factor  in  the  kind  and  quantity  of  food  of  birds..  487 

Solved  and  unsolved  problems  in  economic  ornithology  488 

Summary    490 

Bibliography    494 

Explanation  of  plates  504 


PREFACE 

The  impetus  given  in  late  years  to  the  study  of  the  relations 
of  birds  to  agriculture  in  the  United  States  is  traceable  to  the 
extensive  work  of  the  United  States  Bureau  of  Biological  Survey. 
From  the  organization  of  this  department  of  the  United  States 
Department  of  Agriculture,  July,  1885,  to  December  31,  1911, 
members  of  the  Biological  Survey  have  prepared  and  published 
one  hundred  and  thirty-one  documents  relating  wholly  or  in  part 


1914]   Bryant:  Economic  Status  of  the  Western  Meadowlark     379 

to  the  food  of  birds.  Notes  on  the  economic  status  of  over  four 
hundred  species  of  native  birds  and  of  over  fifty  species  of 
foreign  birds  are  to  be  found  in  these  publications.  In  many 
cases  extended  studies  have  been  made  of  the  food  of  birds  by 
the  examination  of  stomach  contents.  In  no  other  country  has 
economic  ornithology  been  accorded  the  attention  it  has  received 
at  the  hands  of  the  United  States  Department  of  Agriculture. 

In  a  newly  settled  state  like  California,  where  large  tracts 
of  land  are  being  brought  under  cultivation,  disturbances  of  the 
natural  order  of  bird  life  arise  in  two  ways.  First,  the  natural 
food  supply  of  birds  is  destroyed  through  cultivation.  Second, 
a  new  source  of  food  is  often  supplied  by  cultivated  crops. 
Hence  birds  become  of  great  economic  importance.  The  variety 
of  conditions  to  be  found  in  so  large  a  state  as  California  makes 
a  study  of  these  economic  relations  of  birds  complex  and  difficult. 
In  spite  of  the  need,  therefore,  of  a  knowledge  of  the  value  of 
birds,  comparatively  little  work  along  this  line  has  been  done 
in  this  state.  Until  recently  two  bulletins  entitled  "Birds  of 
California  in  Relation  to  the  Fruit  Industry,"  by  F.  E.  L.  Beal, 
published  by  the  U.  S.  Bureau  of  Biological  Survey,  and  a  few 
scattered  notes  in  ornithological  literature  afforded  the  only  pub- 
lished material  on  the  economic  relations  of  birds  in  California. 

Complaints  of  the  depredations  of  birds  in  this  state  have 
been  numerous.  The  injury  to  fruit  caused  by  the  linnet  (Car- 
podacus  mexicanus  frontalis)  is  so  great  that  this  bird  is  branded 
as  a  pest  by  the  fruit-grower.  In  recent  years  grain-growers 
have  complained  of  damage  to  sprouting  grain  caused  by  western 
meadowlarks  (Sturnella  neglect  a).  These  birds  have  been  ac- 
cused, and  rightly  so,  of  boring  down  beside  the  sprout  with 
their  long  bills  and  pulling  off  the  kernel  of  grain.  Ranchers 
have  maintained  that  in  some  cases  whole  fields  of  grain  have 
had  to  be  reseeded  because  of  the  loss  occasioned  by  these  birds. 
In  fact,  there  has  developed  so  much  sentiment  against  the 
meadowlark  that  there  has  been  a  persistent  attempt  made  at 
each  legislative  session  to  take  protection  from  the  bird. 

As  the  western  meadowlark  is  a  bird  defended  by  many 
because  of  its  insectivorous  habits,  the  agitation  following  the 
complaints  has  afforded  an  exceptional  opportunity  to  deter- 


380          University  of  California  Publications  in  Zoology    [VOL.  11 

mine  scientifically  the  economic  value  of  this  bird  by  a  thorough 
investigation.  Such  an  investigation  has  been  made  possible 
through  the  patronage  of  the  California  State  Fish  and  Game 
Commission,  which  established  a  research  fellowship  in  the  De- 
partment of  Zoology  of  the  University  of  California.  In  Jan- 
uary, 1911,  I  was  appointed  Fellow  in  Applied  Zoology  on  the 
Fish  and  Game  Commission  Foundation  in  the  University  of 
California,  with  instructions  to  carry  on  an  investigation  into 
the  relation  of  certain  California  birds  to  agriculture.  The  com- 
mission assumed  the  expense  and  through  its  deputies  furnished 
the  material  for  stomach  examination.  The  office  work  and  the 
laboratory  work  have  been  carried  on  in  the  Zoological  Depart- 
ment of  the  University  of  California.  The  investigation  has, 
therefore,  been  conducted  through  the  co-operation  of  the  State 
Fish  and  Game  Commission  with  the  University  of  California. 

To  the  men  past  and  present  who  have  preceded  me  in  this 
line  of  work  and  who  have  furnished  the  world  with  the  under- 
lying facts  which  have  established  the  science  of  economic  orni- 
thology I  wish  to  give  due  credit,  for  without  their  contributions 
this  work  must  necessarily  have  been  far  more  elemental.  The 
helpful  criticisms  and  valuable  suggestions  of  Professor  Charles 
A.  Kofoid  of  the  University  of  California,  under  whose  direction 
the  work  has  been  done,  have  inspired  and  assisted  me  in  the 
task.  To  Dr.  Joseph  Grinnell,  Director  of  the  Museum  of  Verte- 
brate Zoology  in  the  University  of  California,  who  has  often 
given  me  of  his  time  to  discuss  certain  features  of  the  work,  I 
owe  much.  The  help  also  of  Mr.  E.  R.  Ong  as  laboratory  assist- 
ant, and  of  Professor  C.  W.  Woodworth,  Dr.  E.  C.  Van  Dyke, 
Dr.  F.  E.  Blaisdell,  Mr.  John  Bridwell,  Mr.  W.  L.  McAtee,  Miss 
Anna  M.  Lute  and  others  in  the  identification  of  insects  and 
weed  seeds  has  lightened  the  burden  and  facilitated  this  part  of 
the  investigation. 

Although  collections  have  been  made  of  a  number  of  birds 
about  which  complaint  has  been  received  (western  robin,  bicolored 
red-wing,  Brewer  blackbird,  horned  lark,  western  mourning 
dove,  and  roadrunner),  yet,  because  it  was  the  special  object 
of  attack,  effort  has  been  concentrated  on  determining  the  eco- 
nomic status  of  the  western  meadowlark. 


1914]   Bryant:  Economic  Status  of  the  Western  Meadoivlark     381 

The  investigation  has  consisted  primarily  of  field  investiga- 
tion, experimentation,  and  stomach  examination.  A  large  part 
of  the  field  work  has  been  carried  on  at  Lathrop,  San  Joaquin 
County,  California,  a  place  admirably  suited  for  the  work  in 
hand.  Duties  on  the  agricultural  and  horticultural  demonstra- 
tion train,  which  toured  the  state  in  1911  and  1912  under  the 
auspices  of  the  Department  of  Agriculture  of  the  University  of 
California  and  the  Southern  Pacific  Railway  Company,  have 
afforded  additional  opportunity  to  study  conditions  from  one 
end  of  the  state  to  the  other. 

Economic  ornithology  is  a  new  science  and  has  hardly  pro- 
gressed further  than  the  stage  of  preliminary  interest  and  study. 
As  a  result  practically  all  of  the  work  attempted  thus  far 
has  been  of  the  extensive  rather  than  of  the  intensive  sort,  and 
has  been  made  up  largely  of  a  study  of  the  food  of  birds.  In 
this  investigation  the  attempt  has  been  made  to  improve  on  past 
methods  and,  by  determining  the  food  of  birds  taken  in  the  same 
locality  each  month,  or  twice  each  month,  to  furnish  reliable 
evidence  as  to  their  food  throughout  the  whole  year.  A  study 
of  the  bird  in  the  field,  its  depredations,  and  its  life-history,  has 
also  been  made  in  order  that  all  available  evidence  might  be 
obtained. 

Considerable  difficulty  has  been  experienced  in  that  there 
has  been,  and  now  is,  a  difference  of  opinion  as  to  the  criteria 
to  be  used  in  the  determination  of  the  economic  status  of  a  bird. 
The  ideas  which  have  been  advanced  in  the  past,  and  even  those 
of  the  present  day,  appear  to  be  unsatisfactory,  or  at  least  un- 
trustworthy. It  seemed,  therefore,  that  a  review  of  past  methods, 
with  the  addition  of  such  new  ones  as  appeared  to  be  valuable, 
might  prove  not  only  interesting  but  of  considerable  value  to 
future  workers  in  the  field.  A  similar  lack  of  information  re- 
garding methods  of  stomach  examination  has  been  evident.  A 
detailed  account  of  the  method  used  in  this  investigation,  there- 
fore, seems  justified. 

The  service  which  birds  render  to  agriculture  has  doubtless 
been  overemphasized.  On  the  other  hand,  the  position  taken 
by  some  that  birds  are  of  no  value  as  insect  and  weed-seed  de- 
stroyers hardly  seems  justified  by  the  facts.  If  there  be  a  mis- 


382          University  of  California  Publications  in  Zoology    [VOL.  11 

conception  as  to  the  utility  of  birds,  it  is  high  time  we  sought 
to  destroy  it  and  to  establish  truth  in  its  stead. 

The  economic  relations  of  birds  must  necessarily  become  more 
and  more  important.  As  they  do  so,  the  extensive  study  of  the 
past  few  years  will  give  way  to  the  intensive  study  necessary 
to  solve  the  greater  problems  of  the  future.  Probably  no  one 
thing  will  play  a  greater  part  in  the  conservation  of  wild  life 
than  will  this  intensive  study.  Eecognition  of  the  economic  value 
of  a  single  bird  will  stimulate  interest  in  the  protection  of  all. 

This  "work  is  published,  therefore,  in  the  hope  not  only  that 
the  facts  and  data  here  presented  may  be  of  general  interest  and 
of  value  to  future  workers  in  economic  ornithology,  but  that  it 
may  be  a  factor  in  promoting  the  conservation  of  wild  life  in 
California,  a  state  which  still  possesses  enough  of  its  original 
fauna  to  make  its  conservation  important  and  eminently 
desirable. 


INTRODUCTION 

Doubtless  if  our  knowledge  were  not  so  limited  we  might 
be  able  to  find  a  use  for  most  living  things.  As  it  is,  we  designate 
animals  as  useful,  neutral,  or  injurious  because  of  their  effect 
on  ourselves  or  our  interests.  A  thorough  study  of  the  inter- 
relations of  such  animals  often  reverses  our  original  decisions 
regarding  them.  Not  many  years  ago  insects  as  a  class  were 
called  injurious  because  some  of  them  destroyed  certain  crops. 
Today  only  a  part  of  the  insects  are  considered  destructive,  and 
we  are  yearly  finding  that  others  are  of  neutral  or  beneficial 
character. 

Not  many  years  ago  birds  were  looked  upon  either  as  pests 
or  as  marks  for  the  gunner.  Today  most  of  them  are  looked 
upon  as  valuable  assets  of  the  agriculturist.  As  science  in  the 
past  has  slowly  lifted  us  to  a  plane  where  we  study  the  complex 
interrelations  instead  of  the  single  and  obvious  ones,  so  in  the 
future  we  may  expect  that  more  and  more  each  form  of  life  will 
be  found  to  fill  a  particular  niche  in  its  environment  better  than 
any  contiguous  form. 


1914]   Bryant:  Economic  Status  of  the  Western  Meadowlark     383 

And  yet,  viewed  from  the  utilitarian  standpoint,  there  is  a 
certain  value  in  classifying  organisms  as  injurious  or  beneficial. 
The  danger  lies  not  in  the  classification  itself,  but  in  the  risk 
attendant  upon  a  judgment  hastily  made  or  one  based  on  circum- 
stantial or  partial  evidence.  Being  the  dominant  form  of  life 
on  the  earth,  it  is  only  natural  that  we  measure  the  usefulness 
of  things  by  their  immediate  effect  on  ourselves  or  our  interests 
rather  than  on  the  whole  complex  of  nature. 

An  intimate  knowledge  of  the  use  of  wild  life  is  indispensable 
to  sane  conservation.  Anything  known  to  be  useful  may  justly 
demand  protection.  Anything  known  to  be  of  no  utility  in 
nature  may  justly  be  accorded  destruction.  Ignorance  has 
caused  the  waste  characteristic  of  the  past.  Knowledge  must 
prevent  waste  in  the  future. 

What  may  not  seem  to  be  of  use  today  may  be  of  great 
importance  tomorrow.  It  appears  that  the  economic  value  of 
wild  life  seldom  becomes  evident  until  the  form  becomes  extinct, 
or  at  least  diminished  in  numbers.  It  was  only  a  few  years 
ago  that  fish  were  so  abundant  that  no  attention  whatever  was 
paid  to  their  life-histories.  Today  the  study  of  ichthyology, 
including  fish-hatching,  is  a  necessity,  in  order  that  the  supply 
of  this  kind  of  food  may  continue  to  be  available.  Not  many 
years  ago  people  believed  that  there  was  an  inexhaustible  supply 
of  game.  Today  strict  game  laws  and  the  most  careful  conser- 
vation alone  prevent  the  extinction  of  many  forms. 

Fifty  years  ago  the  farmer  in  the  east  may  have  lost  some 
grain  and  corn  from  the  depredations  of  birds,  but  he  either 
planted  an  extra  acre  or  two  to  make  up  for  the  loss  or  took  it 
as  a  matter  of  course.  At  the  present  time,  however,  when  we 
find  not  only  much  of  the  available  land  under  cultivation,  but 
even  that  cut  up  into  small  tracts  and  men  attempting  to  earn 
a  living  on  ten  or  twenty  acres  instead  of  on  eighty  or  a  hundred, 
the  depredations  of  birds  are  more  noticeable.  The  loss  of  a 
sack  of  grain  is  hardly  noticeable  in  a  large  field,  but  let  the 
same  amount  be  lost  in  a  two-  or  three-acre  field  and  the  loss 
becomes  relatively  important  and  very  apparent.  It  is  only 
natural,  therefore,  that  at  the  present  time  complaints  against 
birds  are  more  frequent  and  more  insistent. 


384          University  of  California  Publications  in  Zoology    [VOL.  11 

Depredations  probably  increase  also  as  the  natural  food  sup- 
ply of  birds  is  destroyed  and  they  have  to  rely  on  the  products 
of  civilization.  The  change  of  food  caused  by  change  in  environ- 
ment sometimes  causes  an  increase  in  the  number  of  birds  of  a 
species,  and  thus  increases  the  extent  of  the  depredations.  Ap- 
parently there  are  a  number  of  birds  in  California  which  have 
directly  profited  by  the  change  of  environment  and  are  increasing 
in  numbers.  The  linnet,  western  meadowlark,  and  mockingbird 
undoubtedly  belong  to  this  class. 

As  the  crops  change,  and  consequently  the  food  supply,  we 
may  even  expect  that  in  the  future  the  food  habits  of  birds  will 
change.  Hence  a  knowledge  of  the  food  habits  of  birds  at  the 
present  time  may  be  of  far  greater  value  in  the  future,  when 
such  data  are  needed  for  comparison. 

Whether  it  is  best  to  destroy  certain  birds  because  of  their 
depredations,  or  to  preserve  them  because  of  their  value  as  insect 
or  weed-seed  destroyers,  has  become  a  real  problem.  There  is 
not  a  farmer  who  is  not  at  some  time  of  the  year  confronted 
with  this  problem.  In  a  newly  settled  country  the  question  as 
to  the  value  of  certain  birds  is  often  of  grave  importance.  If 
nature  were  not  so  closely  woven  together  we  might  easily  solve 
the  problem  by  simply  exterminating  those  birds  which  cause 
damage.  In  the  early  days  this  was  tried.  It  is  experience  that 
has  taught  us  the  danger  attendant  on  the  indiscriminate  ex- 
termination of  any  form  of  life. 

The  problem  stated,  the  next  thing  to  be  considered  is :  How 
shall  it  be  solved?  Observation  has  proved  an  unsafe  method  of 
determining  the  true  value  of  a  bird.  Mere  sentiment  fails  to 
convince  a  large  number  of  the  class  of  people  deeply  interested 
from  actual  contact  with  the  problem.  The  method  which  has 
proved  the  most  dependable  is  a  thorough  scientific  investigation. 
The  problem  is  complex.  It  involves  a  knowledge  of  the  life- 
history  of  birds,  insects,  and  plants,  a  conception  of  the  inter- 
action of  organisms,  and  an  appreciation  of  the  accompanying 
ecological  relations. 

A  scientific  investigation  as  a  means  of  determining  the  status 
of  a  bird  presents  just  as  great  possibilities  as  this  method  has 
in  other  fields.  In  preventive  medicine  we  see  the  results  of 


1914]   Bryant:  Economic  Status  of  the  Western  Meadowlark     385 

scientific  method.  The  efficiency  of  the  forest  service  can  be 
attributed  to  the  same  methods.  If,  then,  this  type  of  investi- 
gation can  help  us  to  conserve  our  health  and  our  forests,  it 
should  also  lead  to  the  best  method  of  conserving  our  native 
birds. 

Professor  Charles  S.  Minot  has  denned  the  method  of  science 
as  "the  art  of  making  durable,  trustworthy  records  of  natural 
phenomena. ' '  He  goes  on  to  say :  ' '  The  method  of  science  is 
not  special  or  peculiar  to  it,  but  only  a  perfected  application 
of  our  human  resources  of  observation  and  reflection — to  use 
the  words  of  von  Baer,  the  great  embryologist.  To  secure  relia- 
bility the  method  of  science  is,  first,  to  record,  everything  with 
which  it  deals,  the  phenomena  themselves  and  the  inferences  of 
the  individual  investigators,  and  to  record  both  truly;  second, 
to  verify  and  correlate  the  personal  knowledges  until  they  acquire 
impersonal  validity,  which  means,  in  other  words,  that  the  con- 
clusions approximate  so  closely  to  the  absolute  truth  that  we 
can  be  safety  guided  by  them."  These  statements  justify  the 
use  of  scientific  method  for  any  modern  problem  and  especially 
for  the  problem  in  hand. 

"But  putting  aside  economic  and  utilitarian  considerations, 
there  is  to  some  of  us  a  greater  stimulus  to  solve  the  problems 
of  nature.  With  the  birds,  and  the  insects  and  plants  upon 
which  they  feed,  we  share  a  common  heritage,  and  the  more  we 
learn  of  the  life  of  these,  our  fellow-workers,  the  nearer  we  ap- 
proach solution  of  the  great  riddle  of  the  Universe,  the  mysterious 
law-abiding  scheme  of  Nature.  The  book  of  knowledge  to  which 
we  add  some  iota  is  marred  with  mystery,  superstition  and  error, 
but  each  proved  fact  cleanses  its  pages.  'Facts,'  says  Laing, 
'are  the  spokes  of  the  ladder  by  which  we  climb  from  earth  to 
heaven.'  "  (See  Coward,  1912.) 

The  labor,  time  and  cost  of  such  an  investigation  as  this  is 
amply  justified  by  the  results  to  be  expected.  A  knowledge  of 
the  real  economic  status  of  a  bird  means  dollars  in  the  pocket 
of  the  rancher,  for  the  destruction  of  any  bird  which  causes 
serious  damage,  or  the  preservation  of  any  bird  that  is  a  benefit, 
has  a  direct  bearing  on  the  size  of  the  crop  raised.  Nor  is  the 
value  to  the  rancher  the  only  value  to  be  considered,  for,  as  will 


386          University  of  California  Publications  in  Zoology    [^OL.  11 

be  seen,  a  bird  has  a  certain  value  to  society  that  cannot  be 
reckoned  in  dollars  and  cents. 

Some  entomologists,  seeing  in  insecticides  the  only  successful 
control  measure  against  insects,  are  inclined  to  minimize  the 
value  of  such  a  natural  control  as  birds.  True  it  is  that  birds 
apparently  do  not  prevent  or  entirely  control  insect  outbreaks, 
for  insects  continue  to  ravage  crops,  no  matter  what  the  bird 
population.  However,  if  a  certain  number  of  insects  cause  a 
certain  amount  of  damage,  it  must  follow  that  a  diminution  of 
the  number  of  insects  causing  damage  must  cause  some  dimin- 
ution of  the  damage  done  even  if  it  be  not  proportional.  We  are 
justified  in  saying,  therefore,  that  the  fact  that  birds  destroy 
great  numbers  of  injurious  insects  shows  them  to  be  important 
agents  in  contributing  to  the  safety  not  only  of  crops,  but  of  all 
vegetation.  Judging  from  the  great  numbers  of  insects  destroyed 
at  the  time  of  an  insect  outbreak,  we  can  safely  infer  that  birds 
may  be  instrumental  in  preventing  the  appearance  of  insects  in 
abnormal  numbers,  by  helping  to  keep  the  numbers  near  the 
normal,  which  we  approximate  by  the  phrase,  "the  balance  of 
nature. ' ' 

In  this  practical  age  almost  everything  is  viewed  from  the 
standpoint  of  dollars  and  cents.  Hence  it  is  desirable  that  we 
study  the  economic  value  of  birds.  There  is  danger,  however, 
in  so  doing,  for  such  studies  may  tend  to  minimize  to  a  certain 
degree  a  value  which  cannot  be  expressed  in  dollars  and  cents. 
To  say  that  a  meadowlark  is  worth  so  many  dollars  to  the  rancher 
each  year  may  obscure  its  esthetic  value. 

The  strongest  opponents  of  the  theory  that  birds  are  bene- 
ficial often  emphasize  the  esthetic  value.  The  following  is  a 
quotation  from  one  of  these  opponents  (Baskett,  1910)  :  "Make 
their  song,  beauty,  grace  and  interesting  habits  a  part  of  our 
culture — and  their  preservation  part  of  our  ethics,  but  do  not 
try  to  foist  them  on  the  farmer  as  an  economic  asset,  for  he 
knows  better  in  many  cases.  If  the  soldier  can  make  better 
marches  under  the  martial  influence  of  the  'spirit-stirring  drum' 
and  *  ear-piercing  fife, '  so  can  the  farmer  gather  inspiration  from 
the  activity  and  cheerfulness  of  birds." 


1914]   Bryant:  Economic  Status  of  the  Western  Meadoivlark     387 

One  need  only  point  to  the  place  which  birds  take  in  art  and 
literature  to  prove  their  esthetic  value.  The  inspiration  for 
some  of  the  finest  paintings,  poems  and  other  pieces  of  literature 
has  come  from  a  knowledge  of  bird  life.  That  exhilaration  and 
inward  joy  awakened  with  an  acquaintanceship  with  birds  has 
a  real  value.  What  mental  pictures  stay  longer  with  us  than 
do  those  gained  first  hand  from  nature?  Those  things  which 
make  the  world  more  beautiful  make  it  more  fit  to  live  in. 

Few  birds  there  are  that  have  a  greater  esthetic  value  than 
does  the  western  meadowlark.  One  reason  for  this  is  that  it  is 
a  conspicuous  bird  and  therefore  known  to  every  one.  Its  song 
has  been  pronounced  far  sweeter  than  that  of  its  eastern  relative. 
Its  plumage  and  general  habits  add  to  its  attractiveness.  A 
bird  associated  with  the  fields  and  plains,  it  adds  great  interest 
to  the  general  loneliness  and  monotony  of  our  great  treeless  areas. 
What  person  traveling  along  a  lonely  country  road  has  not  been 
cheered  by  the  bird  which  stands  bobbing  on  many  a  fence  post 
and  telegraph  pole  and  continually  pours  forth  its  "Eh  heu 
wheel'iky,  wheel 'iky,  wheel'iky"?  (For  the  musical  notation 
of  the  song  of  the  western  meadowlark  see  Allen,  1881.) 

The  educational  value  of  birds  has  more  utilitarian  aspects. 
Birds  teach  a  code  of  ethics  exceeded  only  by  that  of  man  him- 
self. The  fidelity  of  parents  to  each  other  and  to  their  helpless 
young  and  the  industry,  cleanliness,  grace,  and  cheerfulness 
exemplified  by  them  add  much  to  the  finer  ideals  of  life. 

The  educational  and  esthetic  points  of  view  can  even  be 
considered  economically.  To  many,  this  type  of  presentation 
detracts  instead  of  adds.  Nevertheless,  it  is  true  that  the  esthetic 
and  educational  value  of  birds  has  its  economic  relations.  Many 
a  summer  resort  is  chosen  because  of  the  abundance  of  birds  in 
the  vicinity,  and  many  a  summer  vacationist  is  influenced  in  his 
choice  of  destination  by  the  presence  of  birds  in  the  vicinity. 
The  value  of  suburban  property  is  enhanced  by  the  presence  of 
birds.  That  many  a  business  man  has  been  attracted  to  certain 
suburban  localities  because  of  the  presence  of  the  meadowlark 
and  its  song  is  self-evident. 

Carrying  this  point  of  view  to  an  extreme  has  often  antag- 
onized certain  classes,  and  herein  lies  a  danger.  Yet  the  facts 


388          University  of  California  Publications  in  Zoology    [VOL.  1.1 

here  presented  can  easily  be  verified.  A  determination  of  the 
status  of  a  bird  must  include  a  study  of  the  bird  from  every 
point  of  view.  To  many  city  folk  the  esthetic  and  educational 
value  of  a  bird  is  the  more  important,  for  they  never  see  it  from 
any  other  point  of  view.  Perhaps  the  rancher  is  an  extremist 
on  one  side  and  the  city  resident  on  the  other.  A  modification 
of  the  views  on  both  sides  is  very  desirable  to  a  sane  appreciation 
of  the  value  of  birds  in  general  and  the  western  meadowlark  in 
particular. 

The  advance  made  in  investigations  of  the  economic  relations 
of  birds  since  Professor  Aughey  (1878)  studied  the  relation  of 
birds  to  the  locust  ravages  in  Nebraska  up  to  the  present,  when 
government  experts  give  the  whole  of  their  time  to  such  inquiries, 
demonstrates  the  growth  of  the  science  of  economic  ornithology. 

A  study  of  this  advance,  however,  shows  that  only  a  beginning 
has  been  made.  Although  we  know  in  general  the  food  habits 
of  our  common  birds,  yet  conditions  vary  so  greatly  that  we 
cannot  definitely  predict  the  food  in  any  given  locality.  The 
work  thus  far  has  afforded  us  a  general  survey  of  the  food  habits 
of  birds  and  in  some  few  instances  has  given  us  definite  knowl- 
edge as  to  the  usual  food  of  certain  birds.  The  thing  that  eco- 
nomic ornithology  has  not  afforded  us  as  yet  is  a  detailed  study 
of  the  food  of  a  particular  bird  in  a  given  locality  throughout  the 
whole  year. 

The  importance,  then,  of  a  thorough  knowledge  of  the  eco- 
nomic relations  of  a  bird  in  addition  to  its  life-history  is  evident. 
It  has  been  left  to  one  of  the  new  sciences,  economic  ornithology, 
to  tell  us  of  these  economic  relations  and  to  explain  the  real  status 
of  birds.  The  agitation  coincident  with  the  establishment  of 
this  science  had  made  known  at  least  five  facts : 

1.  Birds  are  very  largely  insectivorous,  and  as  a  result  are 
important  in  keeping  the  numbers  of  insects  in  check. 

2.  The  amount  of  food  required  by  birds  is  enormous. 

3.  Birds  often  considered  injurious  are  really  beneficial,  and 
vice  versa. 

4.  Birds  change  their  food  habits  and  feed  on  the  kind  of 
food  most  easily  obtained. 


1914]   Bryant:  Economic  Status  of  the  Western  Meadowlark     389 

5.  Birds  are  very  important  in  preserving  that  balance  of 
nature  most  suited  to  the  interests  of  man,  and  their  place  can- 
not be  filled  by  any  other  class  of  living  things. 

These  facts  are  now  familiar.  They  have  furnished  a  basis 
for  a  sane  protection,  have  demonstrated  the  intricacy  of  the 
interactions  of  organisms,  and  have  helped  develop  the  economic 
view  of  birds. 

Although  economic  ornithology  is  fundamentally  the  study 
of  birds  from  the  standpoint  of  dollars  and  cents,  and,  therefore, 
includes  their  use  as  food,  as  cage  birds,  etc.,  yet  emphasis  has 
rightly  been  placed  on  the  study  of  the  food  of  birds.  As  a 
result,  economic  ornithology  is  most  often  used  in  a  restricted 
sense  and  has  reference  to  the  study  of  the  food  of  birds.  Great 
activity  is  evidenced  in  this  line  of  work  at  the  present.  Not 
a  month  passes  that  there  is  not  some  important  contribution  to 
economic  ornithology,  and  there  is  scarcely  an  entomological  re- 
port that  does  not  mention  the  value  of  birds  as  insect  destroyers. 
To  appreciate  the  work  of  the  present,  however,  there  must  needs 
be  some  knowledge  of  the  work  of  the  past.  A  brief  historical 
review  of  the  subject,  with  emphasis  on  the  methods  used,  will 
furnish  this  needed  information. 


HISTORY  OF  METHODS  IN  ECONOMIC  ORNITHOLOGY 

We  need  only  to  examine  in  detail  the  progress  of  our  sciences 
to  be  convinced  that  there  is  such  a  thing  as  evolution.  As  we 
interpret  their  progress  step  by  step,  and  thus  survey  their  gen- 
eral trend,  it  would  seem  that  the  development  has  been  of  the 
orthogenetic  type.  The  biological  sciences  have  been  a  little 
slower  than  others  in  their  development,  but  they  are  now  taking 
front  rank.  One  of  the  most  marked  tendencies  to  be  noted  in 
history  is  that  of  a  change  from  the  period  when  biologists  drew 
conclusions  from  facts  gained  from  observation  only  to  the 
present  period  when  more  intensive  study  and  experimental 
evidence  are  demanded. 

The  period  of  time  previous  to  1850  may  be  termed  the  prim- 
itive period,  for  during  this  time  we  find  only  an  occasional 
mention  of  the  food  habits  of  birds,  the  entire  time  of  the  workers 


390          University  of  California  Publications  in  Zoology    [VOL.  11 

in  this  field  having  been  given  over  to  classification  and  habit 
notes.  The  following  period  (1850-1865)  marks  the  time  when 
the  specialized  science  of  economic  ornithology  was  founded.  It 
was  a  period  of  interest  and  agitation.  Before  the  conclusion  of 
this  period  the  modern  methods  of  investigation  were  introduced 
into  America  by  Jenks  (1860).  The  period  since  1865  is  best 
considered  the  modern  period — a  time  when  the  attempt  to  reach 
truth  is  backed  by  experimental  evidence  and  the  work  becomes 
intensive  rather  than  general.  (See  Locy,  1908.) 

To  the  first  period  belong  such  men  as  Catesby,  Edwards, 
Forster,  Latham,  Bartram,  Hearne,  and  Barton — men  who  took 
an  active  interest  in  natural  history  and  enriched  ornithological 
literature  with  what  observational  facts  they  were  able  to  glean. 
In  the  latter  part  of  this  first  epoch  there  began  a  marked  tend- 
ency to  gain  more  than  superficial  facts  by  observation,  and  so 
in  the  writings  of  Wilson,  Audubon,  and  Baird  we  find  mention 
of  the  food  of  birds.  (See  Palmer,  1899.) 

It  was  not  till  1860  that  Jenks  (1860)  applied  scientific 
method  to  the  study  of  the  food  of  birds.  Previous  to  this  time 
there  had  been  considerable  agitation  concerning  the  value  of 
birds,  and  many  papers  dealing  with  the  question  appeared  in 
agricultural  journals.  Le  Baron's  "Observations  of  the  Birds 
of  Illinois  Interesting  to  the  Agriculturist"  is  a  good  type. 
Other  writers  at  this  time  were  "Walford,  Holmes,  Kirkpatrick, 
Dodge,  Allen,  Elliott,  and  Samuels.  The  hour  was  ripe,  there- 
fore, for  economic  ornithology  really  to  take  a  place  among  the 
sciences  as  distinct  from  ornithology  itself.  The  work  carried 
on  by  Jenks  (1860),  Treadwell  (1859),  Aughey  (1878),  and 
Forbes  (1880,  1882,  1883,  1903)  gave  the  science  its  real  foun- 
dation, and  inaugurated  the  modern  methods  now  well  exemplified 
and  used  by  the  United  States  Bureau  of  Biological  Survey. 

At  about  the  same  time  we  find  the  science  getting  a  start 
in  Europe.  Jenks  (1860),  when  introducing  the  method  of 
stomach  examination  into  the  United  States,  followed  Prevost's 
(1858)  method.  M.  Florent  Prevost  was  evidently  the  pioneer 
in  Europe.  His  paper,  "A  Memoir  on  the  Alimentary  Regimen 
of  Birds,"  presented  to  the  Imperial  Zoological  Society  of  Paris 
in  the  year  1858,  and  translated  by  Jenks  in  1859,  should  still 


1914]   Bryant:  Economic  Status  of  the  WesternMeadowlark     391 

be  considered  a  classic.  In  it  he  presents  an  original  method 
of  stomach  examination  and  draws  some  very  sane  conclusions 
from  the  results  of  thirty  years'  work. 

It  is  interesting  to  note  that  this  early  worker  had  a  vision 
of  the  very  method  which  is  advocated  at  the  present  time,  but 
which  has  seldom  been  followed  in  detail.  He  says :  "It  ap- 
peared that  it  would  be  of  interest  to  gather,  at  different  periods 
of  the  year,  the  stomachs  of  all  birds  which  it  might  be  possible 
to  procure,  to  examine  the  contents,  to  note  down  the  exact  results 
of  this  examination,  with  the  date  of  the  observation,  and  to 
preserve  these  pieces  in  order  to  form,  in  time,  a  collection  by 
means  of  which  one  can  in  the  future  verify  each  of  the  regis- 
tered facts. ' '  He  goes  on  to  point  out  what  he  had  accomplished 
in  thirty  years'  work  and  the  methods  which  he  used  in  pre- 
serving stomach  contents.  These  methods  were:  drying  and 
mounting  on  cards,  drying  and  preserving  in  a  vial,  and  pre- 
serving in  alcohol. 

His  method  of  examination  appears  to  have  been  thorough, 
for  he  suggests  that  an  "attentive  examination,"  in  many  cases, 
made  "fragments  such  as  antennae,  jaws,  lips  with  their  feelers, 
feet,  and  often  entire  heads"  give  the  means  of  determining  the 
family,  genus,  and,  in  some  cases,  even  the  species.  Following 
this  is  a  discussion  of  the  results  of  the  work.  The  tables  used 
are  described  thus :  "To  this  end  I  have  drawn  up  a  uniform 
table  for  all  the  species  of  birds ;  each  copy  of  this  table  concerns 
a  species  whose  name  figures  at  the  head.  It  represents  a  series 
of  columns,  of  which  each  bears  the  title  of  an  alimentary  regi- 
men ;  it  is  in  these  columns,  and  conformably  to  their  title,  that 
I  have  inscribed  both  the  date  of  the  observation  and  the  indi- 
cation of  the  objects  found  in  the  stomachs.  In  fine,  each  of 
these  tables  contains  a  sufficient  number  of  lines  to  register 
observations  made  during  twelve  months  of  the  year,  and  at  five 
different  dates  in  each  month." 

Early  workers  in  America  have  often  failed  to  consider  the 
food  of  a  bird  for  the  whole  year.  They  have  also  failed  to  take 
into  consideration  the  fact  suggested  by  Prevost  in  the  following 
words:  "The  studies  which  I  have  pursued  after  the  method 
indicated  above  will  establish  the  fact  that  the  same  species  of 


392          University  of  California  Publications  in  Zoology    [VOL.  11 

bird  changes  its  food  according  to  the  age  and  season  of  the 
year."  Also  that  "the  moment  when  certain  insects  inundate 
a  country  with  individuals  without  number,  .  .  .  coetaneously, 
this  very  abundance  seems  to  invite  a  crowd  of  different  species 
of  animals  to  feed  upon  them."  It  was  not  till  1882-1885,  nearly 
twenty-five  years  later,  that  Forbes  clearly  pointed  out  these 
interrelations  between  birds  and  insects. 

Prevost  concludes:  "I  am  in  the  course  of  proving  that 
birds  are  in  general  much  more  useful  than  injurious  to  our 
crops,  and  that  even  in  respect  to  the  greatest  part  of  the  graniv- 
orous  species  the  evil  which  is  done  to  us  at  certain  times  is 
largely  compensated  by  the  destruction  of  insects  which  they 
accomplish  at  other  times.  It  is  important,  then,  that  we  do 
not  destroy  these  species,  but  only  divert  them  from  the  crops 
when  they  injure  them.  Their  destruction  would  permit,  with- 
out counterbalance,  the  development  of  many  species  of  insects 
more  fatal  still  to  agriculture.  The  study  of  the  alimentary 
regimen  has  furnished  me  also  some  information  which  I  believe 
useful  in  comprehending  the  reunions,  the  separations,  and  peri- 
odical emigrations  which  are  observed  so  commonly  among  birds." 
(See  Prevost,  1858,  translation  by  J.  W.  P.  Jenks.) 

Since  the  work  of  Prevost,  economic  ornithology  has  grown 
rapidly.  Germany  has  probably  been  most  active  in  the  work. 
Hawks  and  owls  have  received  the  most  attention  throughout 
Europe,  probably  for  two  reasons.  They  have  been  most  widely 
attacked  because  of  their  size,  and  their  value  is  most  apparent 
upon  investigation.  A  number  of  societies  and  institutions  scat- 
tered over  the  continent  are  actively  engaged  in  studying  the 
economic  status  of  birds.  Chief  of  these  are  the  Kaiserliche 
Anstalt  fur  Land-  und  Forstwirtshaft  zu  Berlin,  Ornithologische 
Gesellschaft  in  Bayern,  Paris  Museum  of  Natural  History,  and 
the  Koniglich  Ungarische  Ornithologische  Centrale.  The  names 
of  Berlepsch,  Key,  Ouster,  Rorig  (1903),  and  Hollrung  (1906) 
have  become  well  known  as  workers  in  this  field  in  Germany. 
Rorig  is  the  one  man  who  has  attempted  a  computation  of  the 
comparative  amounts  of  food  by  a  weight  method.  Csiki  (1909) 
and  Greschik  (1910,  1911)  have  been  the  principal  workers  in 
Hungary.  Their  researches  have  been  mainly  confined  to  the 
birds  of  prey. 


1914]   Bryant:  Economic  Status  of  the  Western  Meadoivlark     393 

A  committee  appointed  by  the  British  Association  for  the 
Advancement  of  Science  is  now  investigating  the  feeding  habits 
of  British  birds  by  a  study  of  the  contents  of  the  crops  and 
gizzards  of  both  adults  and  nestlings,  and  by  collation  of  obser- 
vational evidence,  with  the  object  of  obtaining  precise  knowledge 
of  the  economic  status  of  many  of  the  commoner  birds  affecting 
rural  science.  Data  as  to  the  environmental  conditions  under 
which  the  bird  was  feeding  and  the  available  food  supply  are 
obtained  with  each  specimen.  All  data  obtained  from  the  stomach 
examination  are  tabulated,  and  the  weight  of  the  bird  and  the 
condition  and  weight  of  the  gizzard  contents  are  recorded. 

The  United  States  Department  of  Agriculture  has  carried  on 
the  most  extensive  work  in  economic  ornithology  ever  attempted 
by  one  institution.  Study  along  this  line  was  begun  in  1885. 
Since  that  time  over  sixty  thousand  stomachs  of  birds  have  been 
examined,  and  the  results,  with  the  addition  of  data  collected  in 
the  field,  have  been  published  in  more  than  one  hundred  and 
thirty  bulletins. 

Other  investigations  have  also  been  conducted.  The  most 
extensive  work  has  been  done  in  Illinois  by  the  pioneer  economic 
ornithologist,  Professor  S.  A.  Forbes.  Massachusetts,  Wisconsin, 
and  Pennsylvania  have  also  carried  on  investigations,  the  work 
being  done  by  Forbush  (1908),  King  (1883),  and  Warren  (1888) 
respectively.  Practically  every  state  has  been  supplied  with 
some  literature  on  the  subject  by  the  state  university  or  the 
agricultural  experiment  station. 

At  the  third  ornithological  congress  at  Paris  in  1900,  the 
section  of  economic  ornithology  and  bird  preservation  reported 
in  favor  of  urging  all  countries  and  even  their  governments  to 
take  up  seriously  the  subject  of  the  utility  or  harmfulness  of 
birds  as  being  of  the  greatest  economic  importance.  It  was 
urged  that  "inquiries  should  be  instituted  on  regular  business 
lines,  that  migratory  and  non-migratory  species  alike  should  be 
observed  during  every  month  of  the  year  and  for  several  years 
in  succession,  that  the  contents  of  their  stomachs  should  be  care- 
fully noted,  and  lists  prepared  of  their  action  towards  the 
farmer's  crops." 

The  amount  of  careful  work  along  these  lines  which  has  been 
done  since  this  time  is  very  encouraging.  The  last  few  years 


394          University  of  California  Publications  in  Zoology    [VOL.  11 

have  seen  a  number  of  the  leading  magazines  take  up  the  subject 
of  the  economic  value  of  bird  life,  and  with  illustrated  articles 
they  have  brought  to  the  attention  of  many  the  value  of  birds 
to  the  farmer. 

Largely  because  of  their  depredations,  the  demand  for  an 
intimate  knowledge  of  the  food  of  birds  has  become  very  pressing, 
and  yet  it  is  interesting  to  note  that  attention  to  the  economic 
side  of  ornithology  was  not  aroused  by  the  depredations  of  birds, 
but  by  the  depredations  of  insects. 

A  comparison  of  the  methods  used  up  to  the  present  will 
clearly  show  the  progress  which  has  been  made  in  the  science  of 
economic  ornithology.  Such  a  comparison  is  afforded  by  the 
following  sequence  of  methods  used  in  determining  the  economic 
value  of  birds: 

Sequence  of  methods  used  in  determining  the  economic  value 
of  birds : 

1.  Observational  notes  on  the  food  of  birds.     (Wilson,  1808- 
1814;  Audubon,  1827-1838.) 

2.  Critical  observational  study  of  the   food  of  birds.      (Le 
Baron,  1855;  Holmes,  1857;  Weed,  1903.) 

3.  Examination  of  the  stomach  contents  of  birds.      (Jenks, 
(King,  1883.) 

4.  Experimental  feeding  of  captive  birds.    (Treadwell,  1859.) 

5.  Observation  plus  stomach  examination.      (Aughey,   1878; 
Judd,  1902.) 

6.  Observation  plus  stomach  examination  plus  experimenta- 
tion.    (Forbes,  1903 ;  U.  S.  Biological  Survey.) 

Similar  progress  can  be  noted  in  the  methods  used  in  de- 
termining the  food  of  birds.  Their  sequence  has  been  as  follows : 

1.  Investigation  of  food  with  no  reference  to  time  or  locality. 
(King,  1883.) 

2.  Investigation  of  food  at  time  and  locality  of  depredations. 
(Forbes,    1903;    Wilcox,    1892;    Aughey,    1878;    Bryant,    1911, 
1912d.) 

3.  Investigation  of  food  according  to  the  month,  regardless 
of  exact  locality.     (Jenks,  I860;  Beal,  1907,  1910.) 


19M]   Bryant:  Economic  Status  of  the  Western  Meadowlark     395 

4.  Investigation  of  food  according  to  the  month  in  the  same 
locality.     (Forbes,  1903;  Bryant,  1912a.) 

5.  Investigation  of  food  according  to  the  month  in  the  same 
locality,  with  a  comparison  with  many  different  localities.     (See 
p.  454.) 

The  determination  of  the  economic  status  of  birds  has  like- 
wise progressed.  The  sequence  of  the  criteria  used  has  been  as 
follows : 

1.  Inferential  evidence. 

2.  Circumstantial  evidence. 

3.  Number  of  injurious  insects  eaten. 

4.  Proportion  of  percentage  volume  of  injurious,  neutral,  and 
beneficial  insects  and  seeds  destroyed. 

5.  Contrast  of  all  harm  vs.  all  good,  including  knowledge  as 
to  life-history. 

From  these  comparisons  it  can  be  seen  that  great  progress 
has  been  made.  To  infer  that  a  bird  is  injurious  simply  because 
it  is  seen  in  a  grain  field  or  orchard,  or  to  brand  it  as  injurious 
because  of  circumstantial  evidence  in  the  form  of  grain  or  fruit 
found  in  the  stomach,  are  obsolete  methods  today.  Furthermore, 
we  recognize  at  the  present  time  that  a  bird  may  eat  some  bene- 
ficial insects  and  still  be  a  valuable  bird.  Nothing  less  than  a 
knowledge  of  the  food  for  the  whole  year,  combined  with  a 
knowledge  of  the  life-history  of  the  bird  concerned,  allowing  a 
balance  of  all  the  benefits  conferred  with  all  the  damage  done, 
meets  the  requirements  of  the  present. 


INVESTIGATION  OF  THE  ECONOMIC  STATUS  OF  THE 
WESTERN  MEADOWLARK  IN  CALIFORNIA 

Interest  centered  around  the  meadowlark  for  some  time  pre- 
vious to  the  institution  of  an  investigation.  A  rather  dormant 
complaint  against  the  depredations  of  the  meadowlark  in  sprout- 
ing grain  fields  was  brought  to  a  head  in  a  bill  (no.  229)  intro- 
duced by  Assemblyman  Stuckenbruck  of  San  Joaquin  County 


396  University  of  California  Publications  in -Zoology    [VOL.  11 

into  the  State  Legislature  on  January  11,  1909.  The  bill,  which 
proposed  to  amend  section  637  of  the  penal  code  of  California, 
passed  through  the  committee,  but  was  refused  passage,  the  vote 
standing  32  to  28.  On  the  motion  to  reconsider,  the  bill  was 
again  brought  to  a  vote  and  passed  with  a  vote  of  41  to  28.  The 
Committee  on  Fish  and  Game  of  the  Senate  reported  favorably 
on  the  bill,  but  it  was  refused  passage  on  a  vote  of  17  to  12. 

In  1911  Assemblyman  Stuckenbruck,  at  the  request  of  his 
constituents,  introduced  a  similar  bill  with  the  proviso  that  in 
the  counties  of  Tehama,  Butte,  Sutter,  Sacramento,  Yolo,  Colusa, 
Glenn,  San  Joaquin,  Stanislaus,  Tulare,  and  Kings  the  meadow- 
lark  be  not  included  among  the  birds  protected  by  the  act,  hoping 
thus  to  allay  the  opposition  met  from  other  parts  of  the  state  at 
the  former  session  of  the  legislature.  This  bill,  being  referred 
to  the  Committee  on  Fish  and  Game,  was  returned  to  the  Assem- 
bly with  a  majority  report  in  favor  of  its  passage  and  a  minority 
against  its  passage.  It  failed  of  passage  on  March  20. 

Continued  complaints  from  the  farmers  and  fruit  growers 
of  the  state  have  been  made  to  the  State  Fish  and  Game  Com- 
mission regarding  the  losses  to  crops  caused  by  the  depredations 
of  birds.  The  commission  has  been  repeatedly  urged  to  take 
strong  measures  to  avert  the  damage  done.  The  usual  measure 
urged  is  that  the  particular  bird  in  question  should  be  placed 
on  the  unprotected  list.  On  the  other  hand,  many  scientists  and 
others  interested  in  birds  have  pointed  out  the  fact  that  birds 
confer  a  great  benefit  in  keeping  down  the  number  of  injurious 
insects  and  weed  seeds,  and  thus  they  fill  a  niche  in  the  economy 
of  nature  most  suited  to  mankind  which  is  not  and  can  not  be 
filled  by  any  other  form  of  life.  Experience  has  shown  that 
many  belonging  to  the  first  class  have  based  their  complaints  on 
circumstantial  or  partial  evidence  or  on  evidence  not  sufficiently 
reasoned  out.  Furthermore,  these  complaints  have  brought  out 
the  fact  that  really  very  little  is  known  of  the  food  habits  of 
birds  of  California.  Certain  it  is  that  a  knowledge  of  the  food 
habits  of  a  bird  is  necessary  to  a  determination  of  its  economic 
status.  As  a  result,  therefore,  the  commission  thought  it  wise 
that  legislation  should  be  based  on  scientific  investigation  as  to 
the  value  of  birds,  and  not  on  circumstantial  evidence.  Conse- 


1914]   Bryant:  Economic  Status  of  the  Western  Meadowlark     397 

quently  an  investigation  into  the  relations  of  the  birds  of  the 
state  to  agricultural  and  other  interests  was  instituted. 

The  institution  of  the  investigation  was  largely  due  to  the 
interest  and  energy  of  Mr.  John  P.  Babcock,  Chief  Deputy  of  the 
California  State  Fish  and  Game  Commission,  1910-11,  and 
Professor  Charles  A.  Kofoid  of  the  Department  of  Zoology  of 
the  University  of  California. 


A  COMPARISON  OF  METHODS  IN  ECONOMIC 
ORNITHOLOGY 

The  attempt  to  show  the  amount  of  the  different  kinds  of 
food  contained  in  the  stomachs  of  birds  has  led  to  the  use  of 
two  distinct  methods,  both  of  which  must  be  considered  valuable, 
and  both  of  which  approximate  the  end  sought.  A  method 
introduced  by  King  (1883),  and  later  used  by  Newstead  (1908), 
gives  the  total  number  of  birds  taking  the  different  kinds  of 
food  compared  with  the  total  number  of  stomachs  examined. 
The  second  method,  employed  by  the  United  States  Biological 
Survey,  depends  entirely  upon  the  comparative  volume  of  the 
different  kinds  of  food  found  in  the  stomach,  calculated  in  per 
cent  of  total  volume  and  averaged.  A  third  method,  in  which 
actual  counts  of  the  insects  found  are  made,  has  been  used  in  a 
few  instances  (Mason  and  Lefroy,  1912;  Fisher,  1893).  Workers 
in  this  field  in  Great  Britain  have  used  the  numerical  system 
almost  entirely,  depending  for  a  criterion  upon  the  number  of 
birds  taking  a  certain  kind  of  food. 

Since  all  of  these  methods  appear  to  furnish  certain  infor- 
mation not  furnished  by  the  others,  a  combination  of  all  three 
methods  has  been  used  in  this  investigation.  Dependence  is  laid 
on  the  first  method  for  an  idea  of  the  percentage  of  birds  of  a 
species  feeding  on  a  particular  insect,  on  the  second  for  an  idea 
of  the  comparative  amounts  of  the  different  kinds  of  food  taken 
by  individuals  and  by  the  species,  and  on  the  third  for  an  idea 
of  the  actual  numbers  of  the  different  elements  of  food.  The 
counting  of  weed  seeds  and  insects  found  in  the  stomach  of  a 
bird  is  difficult  and  fruitful  of  error.  Yet  the  fact  that  smaller 


398          University  of  California  Publications  in  Zoology    [VOL.  11 

numbers  are  always  counted  than  truly  exist,  owing  to  the  com- 
minuted condition  of  some,  makes  it  evident  that  no  exaggeration 
is  possible  here.  Consequently  it  affords  dependable  evidence  as 
to  the  numbers  of  weed  seeds,  insects,  etc.,  taken  by  a  bird.  On 
the  other  hand,  the  percentage-of-volume  method  can  be  depended 
upon  only  to  furnish  an  idea  of  the  comparative  quantities  of 
the  different  kinds  of  food.  Personal  error  in  estimating  has 
to  be  allowed  for  in  this  method,  for  whereas  a  certain  insect 
might  be  to  the  eye  of  one  person  ten  per  cent  of  the  volume,  it 
might  represent  fifteen  per  cent  to  the  eye  of  another. 

The  furnishing  of  complete  data  as  to  the  bird  whose  stomach 
is  examined  (date,  locality,  kind  of  field,  collector,  etc.)  should 
afford  information,  first,  as  to  the  variation  in  the  amount  of 
food  taken  by  birds  during  the  day,  month,  and  year,  and  second, 
the  food  preference  of  birds  in  a  given  locality  and  in  different 
localities.  The  record  of  the  exact  time  of  day,  the  month,  and 
year  when  the  bird  was  collected  furnishes  the  basis  for  the  first, 
the  record  of  the  habitat,  as,  for  example,  the  kind  of  field, 
orchard,  or  vineyard,  the  basis  for  the  second. 

An  attempt  has  also  been  made  to  improve  the  method  used 
in  determining  the  economic  status  of  a  bird.  As  has  already 
been  pointed  out,  the  economic  status  of  a  bird  was  originally 
determined  by  inference.  A  bird  in  the  grain  field  must  be 
eating  grain  and  therefore  is  injurious.  Experience  has  taught 
that  such  reasoning  is  fraught  with  error.  And  further  exper- 
ience has  taught  us  that  even  though  a  bird  may  cause  consid- 
erable damage,  yet  because  of  its  usefulness  as  a  weed-seed 
destroyer,  as  an  insect  destroyer,  or  as  a  bird  important  in  keep- 
ing the  balance  in  nature  most  suited  to  man,  it  may  be  more 
beneficial  than  harmful.  At  one  time  the  total  good  accom- 
plished by  a  bird  was  held  to  inhere  in  the  number  of  injurious 
insects  it  destroyed.  Today,  although  we  still  retain  this  idea, 
we  see  a  little  further  and  conclude  that  a  bird  may  be  beneficial 
because  it  destroys  insects  (almost  all  insects  being  potentially 
destructive),  and  not  because  it  chooses  a  particular  class  of 
insects  arbitrarily  classified  as  harmful  by  man. 

An  attempt  has  been  made  to  arrive  at  the  average  volume 
of  food  taken  by  the  meadowlark,  by  determining  the  volume 


1914]   Bryant:  Economic  Status  of  the  Western  Meadowlark     399 

of  food  contained  in  a  large  number  of  stomachs  in  cubic  centi- 
meters and  taking  an  average  volume.  This  allowed  the  record- 
ing of  each  stomach  as  being  of  average  volume,  over  the  average, 
or  below  the  average. 

Identification  of  the  various  insects  and  weed  seeds  found 
in  the  stomachs  has  been  difficult.  Help  from  the  experts  of  the 
United  States  Department  of  Agriculture,  from  the  Department 
of  Entomology  of  the  University  of  California,  and  from  others 
has  facilitated  greatly  the  identification.  Certain  field  work, 
embracing  studies  of  the  abundance  of  birds,  the  depredations 
of  birds,  nesting  habits,  the  relation  of  the  birds  to  insect  out- 
breaks, the  kind,  amount,  and  availability  of  food,  and  the  time 
of  digestion,  has  afforded  needed  supplementary  information. 

In  addition  to  the  importance  of  this  investigation  to  agri- 
cultural interests,  it  has  been  fruitful  of  valuable  data  from  the 
standpoint  of  science.  Although  the  investigation  has  been  car- 
ried on  primarily  to  furnish  practical  information  as  to  the  exact 
relation  of  the  western  meadowlark  to  agriculture  and  horticul- 
ture, yet  no  pains  have  been  spared  to  collect  data  of  purely 
scientific  interest.  The  importance  of  a  knowledge  of  life-histories 
has  been  emphasized  only  of  late.  Information  as  to  the  food 
of  any  form  of  life  constitutes  one  of  the  most  far-reaching 
phases  of  its  life-history.  As  Forbes  (1903)  pointed  out: 
"Since  the  struggle  for  existence  is  chiefly  a  struggle  for  sub- 
sistence, a  careful  comparative  account  of  the  food  of  various 
competing  species  and  genera,  at  different  places  and  seasons 
and  at  all  ages  of  the  individual,  such  as  has  not  heretofore  been 
made  for  any  class  of  animals,  cannot  fail  to  throw  much  light 
upon  the  details,  causes  and  effects  of  .this  struggle.  The  flexi- 
bility of  the  food  habits  of  the  widely  ranging  species,  the  direct 
effects  of  normal  departures  from  the  usual  average  of  food 
elements  upon  the  origin  of  variations,  and  the  general  reactions 
of  birds  upon  their  organic  environment,  are  examples  of  subjects 
upon  which  light  should  be  thrown  by  this  investigation." 

In  this  investigation  the  difference  in  food  habits  of  the 
nestling  and  adult  has  been  clearly  demonstrated  by  the  exami- 
nation of  a  large  number  of  specimens.  The  difference  in  the 
kind  and  amount  of  food  taken  by  the  two  sexes  is  made  available 


400  University  of  California  Publications  in  Zoology    [VOL.  11 

for  the  first  time.  Field  investigation  has  been  fruitful  of  an 
increased  knowledge  of  the  general  habits,  nesting  habits,  abun- 
dance, depredations,  and  distribution  of  the  western  meadowlark. 
The  handling  of  so  large  a  number  of  specimens  taken  from 
all  parts  of  the  state  and  during  each  month  of  the  year  has 
furnished  information  as  to  variation,  albinism,  parasitism,  and 
malformation.  Critical  evidence  as  to  the  value  of  certain  so- 
called  protective  adaptations  of  insects  has  also  been  afforded. 
The  investigation  of  the  relation  of  birds  to  insect  outbreaks 
has  emphasized  their  importance  at  such  times  and  furnished 
critical  evidence  as  to  the  interrelations  of  these  organisms. 


THE  WESTERN  MEADOWLARK   (STURNELLA 
NEGLECT  A) 

In  spite  of  its  name,  the  western  meadowlark  is  not  a  true 
lark,  but  belongs  to  the  family  Icteridae  along  with  the  black- 
birds and  orioles.  It  is  easily  recognized  by  its  medium  size, 
gray-  and  brown-streaked  back,  brilliant  yellow  throat,  black 
V-shaped  collar,  and  its  conspicuous  white  outer  rectrices. 

The  meadowlark  is  widely  distributed  over  North  America. 
The  eastern  meadowlark  (Sturnella  magna  magna)  differs  from 
the  one  found  in  the  west  in  size,  color,  and  song.  The  western 
meadowlark  is  slightly  larger  than  the  eastern  bird,  is  paler  in 
color,  and  has  a  much  richer  song.  For  these  reasons  the  western 
form  is  considered  a  distinct  species  and  is  called  the  western 
meadowlark  (Sturnella  neglecta). 

It  is  found  from  Wisconsin,  Illinois,  Iowa,  Texas,  etc.,  west 
to  the  Pacific  Coast,  and  from  central  and  western  Mexico  to 
British  Columbia  and  western  Canada.  It  is  to  be  found  through- 
out the  State  of  California  from  sea  level  to  7000  feet  elevation 
in  the  mountains. 

The  western  meadowlark  is  resident  throughout  the  year.  A 
slight  altitudinal  migration  perhaps  takes  place,  governed  largely 
by  the  available  food  supply,  but  usually  the  bird  is  to  be  found 
in  the  same  general  locality  throughout  the  year. 


1914]   Bryant:  Economic  Status  of  the  Western  Meadowlark     401 

The  western  meadowlark  is  a  conspicuous  bird  of  treeless 
areas  and  a  frequenter  of  meadow,  pasture,  or  uncultivated 
grass  land.  Although  a  poor  flyer  when  compared  to  some  birds, 
the  meadowlark,  with  its  peculiar  hovering  flight,  is  possessed 
of  a  method  of  locomotion  sufficient  for  its  needs.  Its  mode  of 
life  necessitates  but  a  small  amount  of  flying. 

During  the  fall  and  winter  months  meadowlarks  gather  in 
flocks  of  five  to  fifty  or  more.  During  the  spring,  however,  they 
are  seen  singly  or  in  pairs.  Of  a  nervous  temperament,  they 
are  wary  and  do  not  often  allow  of  close  approach.  Both  the 
male  and  female  are  good  singers.  Their  cheerful  and  varied 
song  is  sometimes  given  from  mid-air,  but  more  often  from  a 
fence  post,  shrub,  or  clod. 

The  western  meadowlark  appears  to  be  one  of  the  few  birds 
which  is  profiting  by  the  increased  cultivation  of  land.  Alfalfa 
furnishes  particularly  good  food  and  cover  for  the  bird,  and 
grain  fields  are  often  chosen  for  a  home.  With  the  furnishing 
of  still  more  good  food  and  cover,  combined  with  the  destruction 
of  some  of  its  enemies,  this  bird  may  be  expected  still  further 
to  increase  in  numbers. 

The  western  meadowlark  feeds  almost  exclusively  on  the 
ground.  It  seldom  perches  in  a  tree  of  any  kind.  The  early 
morning  hours  are  spent  in  obtaining  food,  whereas  the  middle 
of  the  day  is  usually  spent  quietly  hiding  in  the  grass. 

The  food,  composed  largely  of  insects,  grain,  and  weed  seeds, 
is  procured  not  only  from  the  top  of  the  ground,  but  also  by 
probing  beneath  the  soil  and  by  searching  under  clods,  manure, 
etc.  Alfalfa  and  grain  fields  appear  to  be  the  favorite  feeding 
grounds  of  these  birds  in  cultivated  districts. 


FIELD  INVESTIGATION 

Next  to  the  knowledge  of  the  food  of  a  bird  in  determining 
its  economic  status  is  a  study  of  the  bird  at  the  scene  of  action, 
or,  in  other  words,  a  knowledge  of  the  habits  of  the  bird.  Evi- 
dence along  this  line  can  be  afforded  only  by  field  investigation. 

Over  a  month's  time  was  spent  at  Lathrop,  San  Joaquin 
County,  California,  studying  the  abundance,  feeding  habits,  nest- 


402  University  of  California  Publications  in  Zoology    [VOL.  11 

ing  habits,  depredations,  etc.,  of  the  western  meadowlark.  As 
has  been  stated,  this  particular  locality  was  chosen  because  it 
afforded  not  only  an  abundance  of  birds,  but  also  a  favorable 
proportion  of  cultivated  and  uncultivated  land,  thus  allowing  a 
study  of  food  preference. 

The  field  work  carried  on  can  be  grouped  under  three  heads : 
studies  of  the  abundance,  of  the  habits,  and  of  the  depredations. 
Studies  of  the  relations  of  birds  to  insects  and  to  insect  outbreaks 
have  also  been  included  in  the  field  investigation,  but  will  be 
discussed  in  another  place.  (See  p.  456.) 


ABUNDANCE  OF  THE  WESTERN  MEADOWLARK 

Several  findings  in  connection  with  the  field  work  have  tended 
to  minimize  somewhat  the  depredations  of  the  meadowlark.  Per- 
haps one  of  the  most  important  is  the  preference  which  the  bird 
shows  for  uncultivated  land.  Censuses  have  absolutely  demon- 
strated that,  during  hours  of  feeding,  western  meadowlarks  are 
more  abundant  in  pasture  land  than  in  cultivated  fields.  Evi- 
dence as  to  the  abundance  of  the  western  meadowlark  is  also 
afforded  by  the  censuses. 

The  following  are  censuses  taken  in  the  vicinity  of  Lathrop, 
San  Joaquin  County,  on  a  two  and  one-half  hour  drive : 

Uncultivated  Cultivated 

( Pasture )         ( Grain  and  alfalfa ) 

February  28,  1912  158  65 

May  31,  1912  69          27 

These  two  censuses  covered  practically  the  same  length  of  time 
and  the  same  territory.  As  the  birds  were  seen  in  flocks  in 
February,  it  is  only  natural  that  more  birds  were  recorded.  May 
being  in  the  nesting  season,  the  birds  were  then  more  widely 
scattered  and  not  so  easily  seen.  It  is  apparent  that  the  meadow- 
lark  prefers  uncultivated  land  even  at  the  time  of  feeding.  A 
similar  census  taken  at  Acampo,  San  Joaquin  County,  during 
an  hour's  walk  (2-3  P.M.),  resulted  as  follows: 

Number 

Orchards,  vineyards,  and  pasture  25 

Grain  fields  ...  2 


1914]   Bryant:  Economic  Status  of  the  Western  Meadowlark     403 

Meadowlarks  were  more  abundant  in  the  vicinity  of  Lathrop 
than  in  the  vicinity  of  Acampo.  There  is  very  little  pasture  land 
in  the  vicinity  of  Acampo.  This,  if  not  the  main  reason,  is  one 
of  the  important  ones  which  account  for  this  contrast  in  abun- 
dance. Another  contrast  in  abundance  can  be  noted  from  the 
following  censuses,  two  taken  at  Los  Banos,  Merced  County, 
July  11,  12,  1912,  and  the  other  at  Merced,  Merced  County, 
July  17,  1912.  Those  at  Los  Banos  were  taken  while  walking 
less  than  five  miles,  and  occupied  about  four  and  three  hours 
respectively.  The  one  at  Merced  was  taken  while  driving  about 
fourteen  miles,  and  occupied  the  time  between  1 :30  and  5 
o'clock. 


Meadow-     Average 

Locality 

Date 

Time 

larks  seen    per  acre 

Los  Banos 

July  11,  1912 

3-4:30 

36             1.0 

Los  Banos 

July  12,  1912 

2-5:15 

67              1.0 

Merced 

July  17,  1912 

1:30:5 

23               .1— 

It  is  evident,  therefore,  that  the  abundance  of  this  species  of 
bird  is  largely  affected  by  locality.  Probably  in  the  last  analysis 
food  supply  is  the  important  factor.  This  brings  us  to  the 
question :  Can  the  western  meadowlark  obtain  its  natural  food 
in  cultivated  fields  as  easily  as  in  the  uncultivated?  If  we  con- 
sider insects  as  vegetable  feeders,  then  we  should  expect  to  find 
the  best  insect  supply  where  plant  growth  was  most  luxuriant. 
The  cultivation  of  land  destroys  much  of  the  natural  plant 
growth,  and  therefore  must  diminish  the  food  supply  of  the 
insects  enough  to  vary  the  abundance.  Grasshoppers,  cutworms, 
and  wireworms  can  usually  be  found  more  abundant  in  grassy 
pasture  land  than  in  orchards  or  grain  fields.  Hence  it  is  a 
natural  consequence  that  we  find  meadowlarks  frequenting  un- 
tilled  land  more  often  than  tilled  land. 

The  censuses  taken  also,  demonstrated  the  fact  that  meadow- 
larks  were  found  in  the  pasture  land  in  greater  abundance  during 
the  middle  of  the  day  than  in  the  morning  hours  of  feeding. 
During  the  hotter  periods  of  the  day  these  birds  hide  in  the 
grass.  Open  fields  are  seldom  chosen  at  this  time  of  the  day. 


404          University  of  California  Publications  in  Zoology    [VOL.  11 

NESTING  HABITS 

The  nesting  season  of  the  western  meadowlark  lasts  from 
March  to  August.  The  nest  is  a  well-concealed  one,  built  of 
dry  grasses  usually  in  grass,  alfalfa,  or  grain  fields,  in  a  de- 
pression in  the  ground.  A  canopy  of  dry  grass  stems  usually 
arches  the  top  of  the  nest  and  a  runway  two  to  five  feet  long 
leads  to  the  nest.  Ofttimes  this  runway  is  the  only  clue  to  the 
location  of  the  nest.  The  female  bird  does  most  of  the  work 
of  incubation  and  feeding  of  the  young,  while  the  male  acts  as 
a  guard.  Eggs  are  usually  five  and  are  white,  variously  marked 
with  brown,  purple,  and  lavender  spots  and  lines. 

Work  in  the  spring  of  1911  and  1912  substantiated  the  fact 
that  western  meadowlarks  usually  nest  twice  each  year.  The 
first  nesting  usually  occurs  in  April  and  May  and  the  second 
in  July  and  August.  Probably  on  an  average  not  more  than 
three  young  are  in  a  brood,  although  the  number  of  eggs  laid  is 
usually  five.  Second  nestings  examined  usually  show  an  incom- 
plete set  of  eggs.  A  preference  for  pasture  land  for  nesting 
sites  was  shown,  at  least  eighty  per  cent  of  the  nests  found  being 
so  situated.  The  time  of  incubation  was  found  to  be  twelve  to 
fourteen  days.  The  young  stay  in  the  nest  but  a  short  time, 
eight  to  ten  days.  Nestlings  are  exposed  to  many  enemies,  such 
as  skunks,  weasels,  rats,  and  hawks,  and  the  number  of  broods 
successfully  reared  is  less  than  that  of  most  other  birds.  That 
over  ten  per  cent  of  the  nests  in  most  localities  are  destroyed  by 
predacious  animals  and  birds  seems  a  very  conservative  estimate. 
These  facts  have  an  important  bearing  on  the  economic  rela- 
tions of  the  meadowlark.  Proximity  of  breeding  grounds  to 
cultivated  crops  naturally  has  an  influence  on  the  amount  of 
damage  done.  The  rate  of  reproduction  influences  the  amount 
of  damage,  owing  to  the  number  of  individuals  to  be  expected 
in  any  locality. 

DEPREDATIONS  OF  THE  WESTERN  MEADOWLARK 

No  small  part  of  the  field  work  has  consisted  in  investigations 
of  the  damage  caused  by  the  western  meadowlark.  In  most  cases 
the  field  work  has  been  supplemented  with  stomach  examinations. 


1914]   Bryant:  Economic  Status  of  the  Western  Meadowlark     405 

The  principal  complaint  lodged  against  the  western  meadow- 
lark  has  been  that  this  bird  destroys  sprouting  grain.  The  field 
investigations  have  proved  that  this  complaint  has  a  real  foun- 
dation. An  interesting  side-light  on  this  habit  is  afforded  by  a 
paragraph  from  Coues's  (1874)  "Birds  of  the  Northwest": 
"In  April,  before  pairing,  hundreds  used  to  frequent  daily  the 
parade  ground  of  Fort  Randall,  where,  as  the  grass  was  yet 
scarcely  sprouted,  good  opportunity  was  offered  of  observing 
their  characteristic  habit — one  not  so  generally  known  as  it  should 
be,  since  it  is  related  to  the  peculiar  shape  of  the  bill.  The  birds 
may  be  seen  scattered  all  over  the  ground,  busily  tugging  at 
something;  and  on  walking  over  the  scene  of  their  operations, 
the  ground,  newly  softened  by  the  spring  thaw,  is  seen  to  be 
riddled  with  thousands  of  little  holes,  which  the  birds  make  in 
search  of  food.  These  holes  are  quite  smooth — not  a  turning 
over  of  the  surface  of  the  ground,  but  a  clean  boring,  like  that 
made  by  sinking  in  the  end  of  a  light  walking  stick;  just  as  if 
the  birds  inserted  the  bill  and  then  worked  it  about  until  the 
hole  was  of  sufficient  size.  Whether  they  bored  at  random,  or 
were  guided  by  some  sense  in  finding  their  prey,  and  what  par- 
ticular objects  they  were  searching  for,  I  did  not  ascertain ;  but 
the  habit  was  so  fixed  and  so  continually  persevered  in  as  to 
attract  general  attention." 

This  habit  of  boring  into  the  ground  to  obtain  sprouting  seeds 
and  possibly  insects  is  therefore  a  habit  of  old  standing,  and  is 
not  one  recently  developed. 

A  careful  investigation  of  a  sprouting  grain  field  where 
meadowlarks  are  abundant  will  demonstrate  to  any  one  that  the 
western  meadowlark  pulls  sprouting  grain.  At  times  the  drill 
row  is  followed  for  distances  of  four  to  six  feet  and  apparently 
every  sprouted  kernel  is  pulled  up  (pi.  21,  fig.  1).  With  its 
long  awl-like  bill,  the  meadowlark  bores  down  beside  the  sprout, 
grips  the  kernel  and  pulls  it  up.  The  kernel  is  occasionally 
eaten,  but  more  often  it  is  simply  crushed  in  the  bill  to  obtain 
the  milk  and  then  dropped  (pi.  21,  fig.  2).  Consequently 
stomach  examination  cannot  be  relied  upon  to  furnish  accurate 
evidence  as  to  the  total  amount  of  grain  thus  destroyed. 

Certain  fields  examined  have  given  evidence  that  the  deeper 
furrows  made  by  drills  wrere  most  frequented  by  the  birds. 


406  University  of  California  Publications  in  Zoology    [VOL.  11 

Whether  this  was  due  to  the  better  cover  or  to  the  ease  with 
which  the  sprouting  grain  could  be  obtained  it  is  impossible  to 
state. 

Where  meadowlarks  are  very  numerous  and  the  field  of  grain 
small  and  isolated,  considerable  damage  results.  In  some  cases 
such  fields  have  had  to  be  resown.  Since  the  loss  of  a  small 
patch  of  grain  means  far  more  to  the  small  farmer  than  does  a 
larger  amount  of  grain  to  the  large  rancher,  the  bird's  depre- 
dations here  are  important.  The  investigations  have  shown, 
however,  that  fields  apparently  badly  damaged  by  meadowlarks 
when  the  grain  was  sprouting  yielded  the  usual  crops  at  harvest 
time.  This  can  be  accounted  for  in  this  way:  Jhe  birds  can 
succeed  in  pulling  the  grain  for  only  a  short  period  of  time  after 
it  appears  above  the  ground.  By  the  time  the  second  and  third 
leaf  appear  the  plant  is  well  enough  rooted  so  that  the  loss  of 
the  kernel,  even  if  it  should  be  removed,  would  not  injure  the 
plant.  Consequently  the  apparent  devastation  is  largely  mini- 
mized by  the  further  sprouting  of  other  kernels  and  the  successful 
survival  of  the  sprouts  of  many  of  the  kernels  removed.  A 
certain  amount  of  thinning  may  at  times  indeed  be  desirable. 
The  lack  of  uniformity  in  the  thinning  accomplished  by  meadow- 
larks  is,  however,  an  argument  against  them. 

Broadcasted  grain,  unless  harrowed  in  very  deeply,  suffers 
more  than  drilled  grain.  It  is  the  universal  verdict  of  the  grain 
ranchers  of  the  state  that  deeply  drilled  grain  suffers  less  than 
the  shallowly  drilled  or  the  broadcasted  grain.  Experiments 
have  shown  that  the  greatest  yield  comes  from  drilled  grain. 

The  following  table  is  from  University  of  California  Publi- 
cations, Agricultural  Experiment  Station,  Bulletin  211: 

KELATIVE  KESULTS  FROM  DRILLED  vs.  BROADCAST  SEEDING  UPON  THE  YIELD 

OF  GRAIN 

Average  of  22  trials 

Barley  Wheat 

Drilled  70.80  34.85 

Broadcast   64.43  31.60 

6.37  bu.        3.25  bu. 

Percentage  increase  9.9  10.3 

Money  value $3.18  $3.12 


1914]   Bryant:  Economic  Status  of  the  Western  Meadowlark     407 

The  proper  depth  for  grain  to  be  planted  in  sandy  soil  is 
three  to  four  inches.  If  all  grain  were  planted  at  this  depth 
very  little  damage  would  be  possible,  for  meadowlarks  are  unable 
to  bore  more  than  two  and  one-half  inches  at  most.  Holes  meas- 
ured average  about  an  inch  and  a  half  in  depth. 

Margins  of  fields  bordering  pasture  land  usually  suffer  most. 
Ofttimes  a  noticeable  difference  in  the  amount  of  grain  growing 
along  the  edges  of  fields  can  be  attributed  to  the  work  of  meadow- 
larks. 

Owing  to  the  concentration  of  large  numbers  of  meadowlarks 
on  a  single  field,  fields  of  grain  planted  early  suffer  most  from 
the  depredations  of  these  birds.  Grain  sown  late  in  the  year 
suffers  much  less,  for. insects  become  available  in  small  numbers 
and  there  is  less  concentration  of  damage,  due  to  the  larger 
amount  of  available  food. 

Meadowlarks  are  more  able  to  obtain  grain  planted  in  sandy 
soil.  Their  ability  to  bore  deeply  into  soil  after  kernels  of  grain 
varies  directly  with  the  hardness  of  the  soil.  Hard,  dry,  adobe 
soil  precludes  attack.  Sandy  soil,  especially  after  being  softened 
by  a  rain,  is  easily  manipulated  to  advantage. 

An  apparent  preference  of  the  birds  for  oats  has  been  shown 
not  only  by  field  investigation,  but  by  the  complaints  of  the 
ranchers  also.  Probably  its  availability  is  a  greater  factor  than 
any  preference  shown  by  the  bird.  That  less  damage  is  possible 
to  wheat  and  barley  because  the  kernel  is  more  easily  removed 
without  damage  to  the  plant  is  one  theory  proposed.  Certain 
it  is  that  there  is  a  difference  in  the  damage  to  adjoining  fields 
of  oats  and  barley. 

Beyond  an  occasional  instance  of  meadowlarks  pulling  sprout- 
ing garden  seeds,  the  only  other  complaint  of  importance  is  that 
they  destroy  melons  by  boring  holes  in  them.  Most  of  the  com- 
plaint has  come  from  the  San  Joaquin  Valley,  and  especially 
from  Delano  and  Pixley,  Kern  County.  Melon  growers,  although 
admitting  that  meadowlarks  cause  considerable  damage  to  melons, 
have  been  unable  to  demonstrate  the  actual  damage  in  the  field. 
Opinion  is  divided  as  to  whether  they  cause  any  damage.  Some 
prominent  growers  affirm  they  are  never  troubled.  Others  com- 
plain of  a  considerable  loss.  For  instance,  the  names  of  two 


408  University  of  California  Publications  in  Zoology    [VOL.  11 

melon  growers  in  the  vicinity  of  Dinuba,  Tulare  County,  were 
handed  in  as  those  of  men  who  were  greatly  troubled.  One  re- 
turned a  verdict  of  "not  guilty,"  and  the  other  reported  that 
the  damage  was  not  very  great.  All  of  the  growers  report  that 
as  soon  as  there  are  broken  melons  in  the  field  the  birds  cease 
to  be  troublesome.  Apparently  the  depredations  of  meadowlarks 
on  melons  have  been  exaggerated. 

There  is  evidence  to  support  the  view  that  meadowlarks  bore 
into  the  melons  to  obtain  water.  Whether  or  not  they  are  at- 
tracted by  the  sweet  taste  we  cannot  say.  The  placing  of  water 
in  a  field  as  an  experiment  would  doubtless  confirm  or  disprove 
this  view. 

An  occasional  complaint  that  meadowlarks  are  injurious  to 
grapes  has  been  received.  Inquiry  in  grape-growing  sections  of 
the  state  has  led  to  the  conclusion  that  such  damage  is  negligible. 
A  number  of  birds  are  destructive  to  grapes,  chief  of  which  are 
the  oriole  and  grosbeak.  Both  of  these  birds  are  well  known  as 
fruit  eaters.  The  meadowlark,  on  the  other  hand,  seldom  turns 
its  attention  to  fruit  of  any  kind.  No  damage  to  grapes  caused 
by  the  meadowlark  has  been  noted  in  the  field. 

Investigations  of  the  damage  caused  by  meadowlarks  has  led 
to  the  following  conclusions: 

1.  The  western  meadowlark  is  destructive  in  sprouting  grain 
fields,  because  of  its  habit  of  drilling  down  beside  the  sprout 
and  pulling  up  the  kernel.  The  amount  of  damage  done  is  de- 
pendent on  the  particular  location,  the  abundance  of  the  birds, 
the  character  of  the  soil,  the  time  of  year,  depth  and  method  of 
planting,  and  the  kind  of  grain.  The  damage  to  oats  is  greatest, 
wheat  suffers  considerable  damage,  whereas  barley  suffers  but 
little.  Broadcasted  grain  suffers  more  than  drilled,  because  not 
being  sowed  so  deeply  it  is  more  easily  obtained  by  meadowlarks. 
The  birds  often  follow  the  drill  row  and  pull  almost  every  kernel. 
Occasionally,  where  meadowlarks  are  very  numerous  and  the 
quantity  of  grain  small,  fields  have  had  to  be  resown.  The  real 
amount  of  damage  done  has  evidently  been  overestimated,  for 
fields  apparently  badly  damaged  have  given  the  average  yield 
later  in  the  year.  After  the  second  and  third  leaf  appears  on 
the  grain,  the  bird  can  do  little  damage.  This  fact  reduces  the 


1914]   Bryant:  Economic  Status  of  the  Western  Meadoivlark     409 

duration  of  their  depredations  to  less  than  two  weeks,  and  conse- 
quently minimizes  the  amount  of  destruction  possible.  Deep 
planting  and  drilling  as  against  broadcasting  are  important  as 
measures  for  protecting  crops. 

2.  Damage  to  other  cereals,  such  as  corn  and  maize,  and  to 
fruit  is  negligible. 

3.  Investigation  of  complaints  that  meadowlarks  are  destruc- 
tive to  melons  has  shown  that  damage  caused  in  this  way  has 
been  exaggerated.     Melon  growers,  although  claiming  that  the 
birds  cause   considerable   damage,   have   often   been  unable   to 
demonstrate  the  actual  damage  in  the  field. 

4.  Censuses  have  demonstrated  that  the  western  meadowlark 
prefers  grass  land  to  cultivated  land,  nearly  forty  per  cent  more 
birds  being  found  in  the  former. 


EXPERIMENTATION  ON  CAPTIVE  BIRDS 

Experimentation  on  captive  birds  as  a  means  of  determining 
food  preference  has  been  suggested  by  Forbes  (1903)  and  Judd 
(1901).  No  doubt  such  experimentation  furnishes  considerable 
evidence  as  to  the  food  preference  of  the  bird  if  carried  on  with 
proper  controls.  Thus  far  this  sort  of  experimentation  has  not 
furnished  dependable  generalization  as  to  what  the  bird  would 
have  taken  under  natural  conditions  (McAtee,  1912).  This  does 
not  mean  that  better  devised  and  controlled  experiments  would 
not  furnish  dependable  evidence. 

The  difficulty  of  keeping  in  cages  birds  with  the  temperament 
of  the  western  meadowlark  and  the  difficulty  of  procuring  for 
them  proper  food  has  prevented  the  use  of  feeding  experiments 
in  this  investigation. 

In  order  properly  to  estimate  the  quantity  of  food  consumed 
daily  it  has  been  necessary  to  determine  the  time  of  digestion. 
It  was  in  this  determination,  and  in  the  determination  of  the 
quantity  of  food,  that  experiments  on  captive  birds  became  of 
value. 

There  are  four  methods  of  determining  the  quantity  of  food 
required  by  young  birds.  First,  the  quantity  of  food  carried 


410          University  of  California  Publications  in  Zoology    IT°L-  U 

to  the  young  by  the  parents  may  be  observed ;  second,  the  stomach 
contents  may  be  examined  and  the  quantity  estimated;  third, 
experimental  feeding  of  caged  birds  may  be  used;  and  fourth, 
the  quantity  of  food  may  be  determined  by  a  daily  weighing  of 
nestling  birds  and  of  their  excreta. 

The  first  method,  owing  to  the  difficulty  of  observing  in  the 
field  the  feeding  of  the  young  of  so  shy  a  bird  as  the  western 
meadowlark,  has  been  largely  neglected  in  the  interest  of  the 
other  more  practical  methods.  However,  some  observations  as 
to  the  number  of  trips  to  the  nest  made  with  food  have  been  made. 
In  one  instance  a  female  western  meadowlark  carried  food  to 
the  nest  three  times  in  twenty  minutes  (6:15-6:45  A.M.).  As 
the  presence  of  the  observer  caused  some  nervousness  on  the  part 
of  the  parent  birds,  this  cannot  be  considered  the  normal  rate. 
AMOUNT  OF  FOOD  REQUIRED  BY  WESTERN  MEADOWLARKS 
Stomachs  of  nestling  western  meadowlarks  examined  con- 
tained as  high  as  two  grams  of  insect  food.  Maxima  of  seven 
large  cutworms,  of  twelve  grasshoppers  (three-quarters  of  an  inch 
in  length),  and  of  eight  beetles  have  been  found  in  the  stomachs 
of  nestlings.  One  stomach  contained  twenty-four  ants  and  parts 
of  a  ground  beetle.  The  volume  of  nestling  stomachs  and  of 
their  capacity  in  terms  of  the  common  elements  of  food  follows : 

Volume  of  average  cutworm  5  c.c. 

Volume  of  average  ground  beetle  3 

Volume  of  average  grasshopper  75 

Volume  of  average  stomach  of  $  western  meadowlark  3.00 

Volume  of  average  stomach  of  $  western  meadowlark  2.50 

Capacity  of  average  <$  stomach  in  cutworms  6 

Capacity  of  average  $  stomach  in  cutworms  5 

Capacity  of  average  £  stomach  in  ground  beetles  10 

Capacity  of  average  $  stomach  in  ground  beetles  8 

Capacity  of  average  $  stomach  in  grasshoppers  4 

Capacity  of  average  $  stomach  in  grasshoppers  3 

A  nestling  western  meadowlark  after  obtaining  no  food  for 
three  hours  was  fed  twenty-eight  small  grasshoppers  (one-half 
inch  in  length),  equal  in  volume  to  about  three  cubic  centimeters. 
Another  one  was  fed  four  grasshoppers  (one  inch  in  length), 
twelve  small  grasshoppers  (one-half  inch  in  length),  one  robber 


1914]   Bryant :  Economic  Status  of  the  Western  Meadowlark     411 

fly,  one  beetle,  and  five  ants.    A  third  one  was  fed  thirty  grains 
of  wheat  inside  of  ten  minutes. 

Weighings  of  nestling  birds  demonstrated  the  fact  that  they 
gained  very  nearly  seven  grams  (0.6  ounce)  in  weight  daily. 
Solid  excreta  averages  0.48  gram.  The  weight  of  excreta  voided 
'  during  twenty-four  hours  must  be  near  3.6  grams,  thus  making 
the  weight  of  food  required  daily  over  ten  grams.  This  com- 
putation does  not  take  into  account  the  weight  of  excretory 
products  given  off  through  the  skin  or  the  weight  of  carbon 
dioxide  given  off  through  respiration.  The  ratios  of  solid,  liquid, 
and  gaseous  excreta  are  not  known.  The  tabular  results  of 
weighings  follow: 

WEIGHINGS  OF  NESTLING  MEADOWLAKKS 

Average  weight  of  2  western 
Time  of  weighing  meadowlark  nestlings 

Nest  No.  1  Nest  No.  2^ 

.75  oz. 
1.25 
1.50 

1.75         .75  oz.       .50  oz. 
2.9         1.00  .75 

2.0 
2.25       1.50  1.25 

1.75  1.50 

Weight  of  egg  ready  to  hatch 135  oz. 

Weight  of  day-old  nestling  25 

Weight  of  eight-day-old  nestling  2.50 

Weight  of  average  adult    4.00 

Although  more  experiments  are  necessary  to  establish  the 
exact  gain  in  weight  of  nestling  birds,  yet  these  experiments  have 
furnished  evidence  as  to  the  enormous  quantities  of  food  con- 
sumed by  nestling  birds.  When  one  considers  that  there  is  a 
gain  of  about  three  ounces  (93.3  grams)  in  weight  inside  of  two 
weeks,  and  that  this  added  weight  must  be  calculated  by  the 
weight  of  food  consumed  minus  the  waste  thrown  off  in  the 
various  forms  of  excreta  and  expired  air,  the  quantity  of  food 
necessary  is  evident.  Not  only  is  enough  food  needed  to  maintain 
energy,  but  an  additional  amount  to  maintain  weight  increment 
is  demanded  during  the  period  of  growth.  Probably  each  young 


Date 
May  26 

Nest  No.  1 
6:00  a.m. 

Nest  No.  2 

May  27 

8:30  a.m. 

May  28 

8:40  a.m. 

May  29 
May  30 
May  31 

7:15  a.m. 
7:45  a.m. 
4:  25  p.m. 

5:45  p.m. 
5:00  p.m. 

June  1 
June  3 

7:  30  a.m. 

6:30  a.m. 
6:15  a.m. 

412          University  of  California  Publications  in  Zoology    [VOL.  11 

western  meadowlark  consumes  something  like  five  ounces  (155.5 
grams)  of  food  during  the  time  it  remains  in  the  nest.  This 
weight  in  grasshoppers  would  mean  311  individuals  and  in  cut- 
worms 415  individuals. 

The  time  of  digestion  of  the  western  meadowlark  was  deter- 
mined by  feeding  captive  juvenile  birds  and  examining  the  con- 
dition of  the  food  at  intervals  after  feeding.  The  following 
table  gives  a  summary  of  the  results  of  these  experiments : 

TABULAR  EESULTS  OF  EXPERIMENTS  ON  THE  TIME  OF  DIGESTION 

Time 

Time  between 

Exper-       without       feeding  and  Food  given  Condition  of  food 

iment          food  killing  on  examination 

1  3%  hrs.       2  hrs.      4  large  grasshoppers          Finely  comminuted 

(Camnula  pellucida)  and  largely  digested 

12  small  grasshoppers 
1  robber  fly 
1  beetle  (Coniontis  sp.) 
5  ants 

2  5%  3%  20  grasshoppers  About  one-quarter  of 

(Camnula  pellucida)          volume  remained;  all 
10  ants  soft  parts  digested 

3  3^  2  30  kernels  wheat  15  kernels  left 

undigested;  hulls 
still  undigested 

4  3%  2  1  May  beetle  Only  hard  parts 

(Ligyrus  gib~bosus}  left  in  stomach 

1  weevil  (Bhygopsis  sp.)       (heads,  wing-covers 
12  grasshoppers  knee  joints,  etc.) 

(Camnula  pellucida) 

5  4  5?  28  grasshoppers  Stomach  empty 


From  these  data  it  can  be  safely  concluded  that  insects  are 
digested  in  two  to  four  hours  and  that  the  stomach  is  completely 
emptied  every  four  hours.  Beetles  and  ants,  owing  to  the  chi- 
tinous  parts,  remain  longer  in  the  stomach  than  do  grasshoppers. 
Cutworms  doubtless  are  digested  much  more  rapidly.  Grain  is 
more  difficult  to  digest  than  insects  and  remains  in  the  stomach 
longer  (four  to  five  hours). 

These  results  compare  very  favorably  with  the  results  of 
similar  experiments  by  other  investigators.  Experiments  carried 
on  by  Treadwell  (1859),  Forbush  (1907),  and  Weed  and  Dear- 
born (1903)  have  demonstrated  that  birds  have  a  very  rapid 


1914]   Bryant:  Economic  Status  of  the  Western  Meadoivlark     413 

digestion,  most  of  them  requiring  from  one  to  four  hours  only 
to  digest  a  meal.  All  evidence  of  food  had  disappeared  from 
the  excreta  of  a  crow  within  two  and  one-half  hours  after  feeding. 
Professor  Treadwell  showed  that  juvenile  robins  digested  a  meal 
every  two  to  four  hours. 

All  evidence,  therefore,  points  to  the  fact  that  four  hours 
can  be  considered  a  sufficient  period  of  time  to  assure  the  diges- 
tion of  the  stomach  contents  of  a  western  meadowlark.  The 
contents  of  the  stomach  at  the  time  the  bird  is  killed  must  have 
been  taken  within  four  hours  previous  to  the  collection  of  the 
bird.  The  daily  consumption  must,  therefore,  be  considered 
about  three  times  the  capacity  of  the  bird's  stomach.  As  the 
birds  start  each  day  with  an  empty  stomach  and  with  the  addi- 
tional stimulus  of  hunger,  the  greater  amount  consumed  during 
early  morning  hours  compensates  for  the  smaller  amount  taken 
during  the  middle  of  the  day. 


EXAMINATION  OF  THE  STOMACH  CONTENTS 

COLLECTION  AND  PRESERVATION  OF  MATERIAL 

Birds  in  sufficient  numbers  to  furnish  reliable  data,  collected 
every  two  weeks  during  a  year,  and  from  over  twenty  different 
localities  in  the  state,  have  been  made  available  through  the  co- 
operation of  the  deputies  of  the  Fish  and  Game  Commission. 
Each  bird  was  tagged  with  data  as  to  date,  time  of  day,  locality, 
kind  of  field  or  orchard,  and  collector.  Dependable  data  re- 
garding abundance  of  food  were  not  available  as  a  general  rule. 
The  birds  were  then  preserved  in  formalin.  On  the  arrival  of 
shipments  at  the  laboratory  the  stomach  (gizzard)  was  removed 
and  data  as  to  the  species  and  sex  of  the  different  birds  added. 
The  tag  bearing  complete  data  was  then  wrapped  with  the 
stomach  in  a  small  cloth,  and  preserved  in  ten  per  cent  formalin 
until  microscopically  examined. 

In  determining  the  localities  where  collections  have  been  made 
an  attempt  was  made  to  select  well-settled  parts  of  the  state 
representative  of  the  different  agricultural  sections.  The  fol- 
lowing instructions  were  sent  to  each  deputy: 


414          University  of  California  Publications  in  Zoology    [VOL.  11 

INSTRUCTIONS   TO   COLLECT   BIRDS   FOR   SCIENTIFIC   INVESTI- 
GATION OF  THEIR  RELATION  TO  AGRICULTURE 
DEAR  SIR: — 

In  order  to  obtain  material  for  our  field  investigation  of  the  relations 
of  game  and  other  birds  to  agriculture,  we  propose  to  collect  specimens 
of  all  the  field  and  orchard  birds  in  the  State  other  than  quail,  ducks, 
geese,  and  crows  during  the  first  and  third  week  of  each  month. 

We  want  especially  one-half  dozen  each  of  meadowlarks,  robins,  and 
blackbirds  collected  during  these  weeks  in  grain  fields  and  the  same 
number  in  vineyards  or  orchards  in  sections  where  there  are  both.  We 
want,  also,  two  doves  from  the  same  section  for  each  of  the  two  weeks 
mentioned.  For  the  present  we  will  confine  our  attention  to  meadowlarks, 
horned-larks,  robins,  and  blackbirds.  Where  practicable  obtain  specimens 
of  each  species  of  your  district  from  the  same  field  or  fields  of  a  like 
character  near  by,  because  we  wish  to  show  just  what  they  feed  upon 
throughout  the  year. 

As  each  bird  is  killed  fill  out  one  of  the  string  record  tags  furnished 
you,  being  careful  to  note  the  hour,  date,  character  of  field,  orchard  or 
vineyard,  the  location  and  the  name  of  the  owners  and  the  nearest  post- 
office.  Then  securely  fasten  a  tag  to  each  bird.  In  writing  upon  the  tags 
use  plain  lead  pencils  only,  as  the  preserving  fluid  will  destroy  ink  and 
indelible  pencil  marks. 

After  filling  out  the  tag,  attach  it  to  the  bird  and  then  record  in  a 
notebook  devoted  to  this  work  a  similar  record  to  the  one  on  the  tag. 

Upon  returning  from  the  field  proceed  to  preserve  the  specimens  as 
follows:  From  any  drug  store  purchase  a  40  per  cent  solution  of  formalin 
or  formaldehyde.  Then  place  in  an  ordinary  two-quart  fruit  jar  one-half 
pint  of  the  40  per  cent  formalin  and  then  fill  up  that  jar  with  clear  water 
and  mark  it  "Jar  No.  1."  Then  place  your  specimens  in  another  jar 
(or  jars)  and  fill  it  with  the  liquid  from  "Jar  No.  1,"  and  seal  the  top 
to  prevent  evaporation.  Don't  place  too  many  specimens  in  a  jar;  give 
them  room  enough  to  become  thoroughly  saturated.  Keep  the  specimens 
in  the  solution,  for  at  least  a  week.  See  that  the  tag  of  each  specimen  is 
uninjured. 

At  the  end  of  each  month  take  your  specimens  from  the  solution  and 
wrap  in  a  cloth  wet  from  the  solution  in  "Jar  No.  1"  and  place  in  a  thin 
cracker  box  or  a  tight  wooden  box  and  ship  as  hereafter  directed,  together 
with  a  copy  of  the  record  from  your  notebook. 

The  solution  from  which  you  remove  your  specimens  can  be  used 
several  times,  supplying  the  necessary  additional  solution  from  ' '  Jar 
No.  1." 

Recember  that  birds  have  no  value  unless  each  is  securely  tagged  with 
a  record  of  the  hour,  date,  etc.;  that  our  work  depends  absolutely  upon 
the  accuracy  and  reliability  of  the  record.  Send  separate  bills  covering 
your  purchase  of  shot,  formaline,  jars,  traveling  and  other  expenses  con- 
nected with  this  work  to  this  office.  Do  not  include  them  on  your  regular 
bills. 

The  necessary  permit  to  take  these  birds  throughout  the  year  is  fur- 
nished you  herewith. 


1914]   Bryant:  Economic  Status  of  the  Western  Meadowlark     415 

Inform  your  local  press  and  others  interested  just  what  you  are  doing 
and  the  object  we  have  in  making  this  collection,  to  wit,  investigating 
the  relations  of  our  birds  to  agriculture,  so  that  there  may  be  no  mis- 
understanding. 

Finding  that  it  worked  a  hardship  on  the  deputies  to  spend 
certain  days  of  each  month  collecting  birds,  the  original  instruc- 
tions were  modified  as  follows: 

Our  order  concerning  the  collection  of  non-game  birds,  for  scientific 
investigation,  is  hereby  modified  to  permit  collecting  at  convenient  times 
instead  of  during  the  first  and  third  weeks  of  each  month.  The  collection, 
however,  must  be  distributed  evenly  so  that  nearly  the  same  number  of 
specimens  are  secured  in  each  bi-weekly  period. 

Under  this  order  we  believe  the  deputies  themselves  can,  while  on 
other  duty,  take  all  the  specimens  needed  for  our  work  and  thereby  greatly 
reduce  expense.  In  consequence  we  direct  that  all  special  collectors  be 
dismissed  and  their  permits  and  collecting  material  taken  over  by  the 
regular  deputies  responsible  for  their  engagement. 


MATERIAL 

The  accompanying  map  (fig.  A)  shows  the  localities  in  which 
collections  have  been  made.  The  localities  from  which  complete 
series,  that  is,  birds  collected  each  month  of  the  year,  were  made, 
and  those  localities  from  which  incomplete  series  were  obtained, 
are  both  indicated.  The  attempt  was  made  to  have  a  minimum 
of  six  specimens  collected  each  month.  In  several  instances 
collections  of  a  dozen  birds  each  month  were  obtained. 

In  order  that  the  work  of  1911  might  be  verified,  collections 
were  continued  at  Live  Oak,  Sutter  County ;  Sacramento,  Sacra- 
mento County ;  Newman,  Stanislaus  County ;  and  Salinas,  Monte- 
rey County,  during  1912.  This  afforded  a  comparison  of  the 
food  in  two  successive  years,  and  has  acted  as  a  check  on  the 
results  obtained  the  first  year. 

A  total  of  2070  stomachs  of  western  meadowlarks  has  been 
available  for  examination,  of  which  number  nearly  two  thousand 
have  been  examined  and  the  results  tabulated.  The  largest  col- 
lection available,  composed  of  one  hundred  and  seventy-five  birds, 
was  from  Hanford,  Kings  County.  The  largest  number  available 
taken  in  a  single  month  was  twenty-four. 


416          University  of  California  Publications  in  Zoology    [VOL. 11 


s  § 
2  •« 


31 

r^       Z 


sppg  snoau         ^       LO       10 


UJOO  UI 


uuaq  ui 

ui 
«! 
sptau 


-euiA  ui 


UtBj 


CO(MQOC5 


COCOC<ICX) 
rHiHCMrH 


CO          <M         O5          TH 


W 


f»I 


t     1 


0>    <V 

oo 


g?    03 

^  >» 


§    B. 


g 


1914]   Bryant:  Economic  Status  of  the  Western  Meadowlark     417 


sppg  snoau 

UJOO  UI 

oc 

0 

CO          00          i—l 

^H 

oo 

00 

flyr 

U7 

H 

0 
r-  1 

1C 

:       co 

H 

1 
•3 

usaq  ui 

N  —  Conclu 
jers  taken 

suap.iBS  ui 

c 

^ 

! 

0 
rH 

I1 

sppg 

oc 
oc 

) 

) 

ir 

t> 

CO 

co 

cc 

) 

05 
00 

s 

M 

w 

H 

9jn^sBd  ui 

c<- 

5         IT 

I 

) 

ir 

i—  I          05 

IT 

r— 

) 

< 

0 

rH 

STOMA 

spas.? 
-auiA  ui 

t> 

•          r~ 

t> 

^ 
T- 

:       os 
i              ^ 

(M 
!>• 

BB 

g 
•a 

spaBqojo  ui 

c 
o- 

5          r- 
9 

*        ^ 

C^ 

o 

1 

(M         rH 
l> 

te 

) 

CO 
CO 

SUITABK 

pa^uasaj 

V 

}          Ci 

T 

5         C^ 
H         1C 

i      «: 

5          r- 

>         C 

T- 

)      'T| 
1         O- 

1       cc 

9 

>      t> 

T— 

0 

1        O 

5         CO         O5         C*" 
t- 

G 
3         r- 

5 
H 

o 

^H 

-daa  sqjuora         c<J       rH       <M 


OQ 


l 

8  s 


W      H* 


H      <! 
i-s       r-a'- 


'-Q      S 

O          o 

03  r-H 

kJ          X! 

W         pQ 

O> 


oo  co    |  o 

§ 


•g  a     o 

K  >,      EH 


b   6 

g 


o 


* 


418          University  of  California  Publications  in  Zoology    [VOL.  11 


LEGEND 

O      Lesstfian  six  montkly  collections 
vl7     Sixtoten  monthly  collections 
I        Complete  series  f< 

'*  c 


or  one  year 


omplete  series  for  two  years 


o^- 


^ 


Fig.  A. — Map  of  California  showing  localities  in  which  collections  of 
western  meadowlarks  have  been  made  for  the  purpose  of  stomach  exami- 
nation. A  complete  series  is  a  minimum  of  six  birds  collected  each  month 
in  a  year. 

Table  I,  pages  416-417,  gives  a  summary  of  the  material 
available. 


EXAMINATION  OF  STOMACH  CONTENTS 

On  removing  the  stomach  (gizzard)  and  tag  from  the  cloth, 
the  stomach  was  carefully  cut  open,  the  incision  being  made  with 
a  scissors  along  the  longest  axis  and  through  the  muscles,  starting 
at  the  cardiac  opening,  and  the  contents  emptied  upon  a  glass 


1914]   Bryant:  Economic  Status  of  the  Western  Meadoidark     419 

plate.  Great  care  was  taken  to  see  that  every  bit  of  the  contents 
was  scraped  from  the  walls  of  the  stomach.  A  Zeiss  binocular 
was  used  in  examining  and  determining  the  material  found. 
Where  possible,  counts  were  made  of  all  vegetable  and  animal 
elements  of  food.  The  amount  of  mineral  matter  used  as  grinding 
material  was  computed  by  a  calculation  in  per  cent  of  the  com- 
parative volume. 

The  comparative  volume  of  each  kind  of  food  was  calculated 
in  per  cent  of  total  volume  contained  in  the  stomach  examined. 
In  the  estimation  of  apparent  volume  there  is  always  a  personal 
error.  However,  since  this  method  is  depended  upon  to  furnish 
evidence  as  to  the  comparative  amounts  and  not  actual  amounts, 
the  personal  error  is  largely  distributed  in  the  averages.  Parts 
of  insects  and  weed  seeds  used  in  identification  were  wrapped 
separately  in  small  pieces  of  paper  to  prevent  their  mixing  with 
the  rest  of  the  stomach  contents. 

Where  possible,  the  heads  of  insects  were  used  as  a  safe  cri- 
terion of  the  number  eaten.  In  many  cases  dependence  was 
necessarily  placed  on  an  enumeration  of  the  mandibles.  In  the 
case  of  grasshoppers  the  mandibles  are  probably  retained  in  the 
stomach  longer  than  the  soft  parts,  but  experiment  has  shown 
that  the  stomach  is  completely  emptied  in  four  hours,  so  that 
it  is  necessary  only  to  give  a  long  enough  period  of  digestion 
to  make  such  an  enumeration  dependable.  The  fact  that  the 
mandibles  of  grasshoppers  may  be  found  along  the  entire  length 
of  the  digestive  tract  also  supports  the  view  that  this  criterion 
is  trustworthy.  Beetles,  bugs,  bees,  and  ants  were  readily 
counted,  because  the  heads  and  thoraces  of  these  insects  remain 
long  undigested. 

Partly  digested  grain  and  weed  seeds  were  computed  in  per- 
centage volume,  but  in  addition  the  undigested  kernels  and  seeds 
were  counted. 

Owing  to  the  finely  comminuted  condition  of  the  food  to  be 
found  in  the  intestines,  that  found  in  the  stomach  alone  was 
used  as  evidence  of  the  food  taken.  The  stomach  alone  gives 
the  best  unit  of  volume.  The  consideration  of  the  food  to  be 
found  in  the  intestines  could  at  best  but  show  evidence  as  to 
food  for  a  longer  time  previous  to  the  death  of  the  bird. 


420          University  of  California  Publications  in  Zoology    [VOL.  11 

In  all  cases  the  stomach  contents  were  preserved  in  vials,  so 
that  verification  of  the  results  will  be  possible  at  any  time.  As 
a  rule,  the  contents  have  been  preserved  by  drying,  but  where 
certain  animal  matter  such  as  larvae  was  present  the  material 
has  been  preserved  in  seventy  per  cent  alcohol. 

After  each  examination  the  kind  and  quantity  of  food  was 
recorded  on  a  stomach  blank.  Complete  data  were  recorded  on 
larger  blanks.  Summaries  giving  the  results  of  the  examinations 
of  the  different  collections  were  made  in  the  form  of  tables. 

IDENTIFICATION  OF  STOMACH  CONTENTS 

Collections  of  insects  and  seeds  for  comparison  were  most 
helpful  in  identifying  the  different  insects  and  seeds.  Insects 
and  seeds,  if  in  good  condition,  can  be  determined  at  least  to 
the  genus  by  this  method.  Floating  out  the  wings  of  certain 
insects  in  water  and  the  mounting  of  other  parts  in  some  clearing 
fluid  are  methods  which  have  had  to  be  resorted  to  occasionally. 

In  the  examination  of  a  large  series  of  stomachs  it  is  nearly 
always  possible  to  obtain  a  fairly  good  specimen  of  an  insect 
which  is  commonly  taken  as  food,  for  some  bird  is  usually  found 
which  has  taken  such  an  insect  just  before  being  killed. 

FOOD  OF  THE  WESTERN  MEADOWLARK  IN  CALIFORNIA 

The  food  of  the  western  meadowlark  is  made  up  of  both 
vegetable  and  animal  matter.  The  vegetable  food  is  largely  com- 
posed of  grain  and  seeds.  The  animal  food  is  made  up  largely 
of  insects.  The  accompanying  diagram  (fig.  B)  shows  the  rela- 
tive amounts  of  the  different  kinds  of  food  taken  during  the  year. 
A  discussion  of  each  kind  of  food  is  followed  by  a  statement  of 
the  "amount  destroyed"  and  the  "economic  importance." 

VEGETABLE  FOOD 
Grain 

Grain  forms  the  largest  percentage  (seventy-five  per  cent)  of 
the  vegetable  food  for  the  year  and  makes  up  thirty  and  eight- 
tenths  per  cent  of  the  total  food  for  the  year.  But  a  small  part 
(one  per  cent)  of  that  found  in  the  stomachs  has  been  sprouted. 


1914]   Bryant:  Economic  Status  of  the  Western  Meadowlark     421 

This  was  therefore  in  all  probability  pulled  from  the  seeded  fields. 
Oats  is  the  grain  most  often  taken.  It  is  not  only  preferred,  but 
is  the  most  available.  Much  (about  seventy  per  cent)  of  that 
found  in  the  stomachs  is  wild  oats  (Avena  fatua).  This  oat  is 
so  mixed  with  the  tame  varieties  that  part  of  it  must  be  consid- 
ered a  loss  to  the  rancher,  for  it  makes  good  feed.  Nearly  half 


PROPORTIONS  OF  DIFFERENT  KINDS  OF 
FOOD    FOR    THE    YEAR 


ANIMAL   FOOD 
VEGETABLE    FOOD 


63.3% 


36.7% 


Fig.  B. — Diagram  showing  relative  amounts  of  different  kinds  of  food 
taken  during  the  year  by  western  meadowlarks. 


422  University  of  California  Publications  in  Zoology    [VOL.  H 

of  all  the  grain  taken  by  the  birds  examined  was  consumed  during 
the  three  winter  months — November,  December,  and  January. 
It  is  apparent,  therefore,  that  availability,  lack  of  insect  food, 
and  possibly  the  sprouting  condition  of  the  grain  are  responsible 
for  this.  Owing  probably  to  the  feeding  habits  of  the  bird, 
mixtures  of  different  grains  are  seldom  found.  The  stomachs 
usually  contained  one  kind  of  grain  only. 

Barley,  because  of  its  greater  availability,  is  more  often  taken 
than  wheat.  The  barbs  are  seldom  found  in  the  stomach.  One 
hundred  and  fifty-five  out  of  one  thousand  and  nine  hundred 
birds  had  eaten  barley. 

Wheat  is  taken  less  often,  probably  because  it  is  less  available. 
One  would  suppose  that  a  grain  without  the  hull  would  be  the 
more  palatable.  Field  corn  (Zea  mays)  was  taken  by  only  seven 
birds  out  of  one  thousand  nine  hundred  and  twenty,  while  ten 
had  eaten  white  milo  maize  (Andropogon  sorghum)  and  other 
varieties  of  Egyptian  corn. 

Quantity  destroyed. — During  the  months  of  December  and 
January  the  meadowlark  feeds  largely  on  grain.  Grain  forms 
nearly  thirty-one  per  cent  of  the  food  for  the  year.  Stomachs 
are  often  found  entirely  filled  with  oats,  barley,  or  wheat.  As 
many  as  thirty  kernels,  with  enough  hulls  to  account  for  as  many 
more,  have  been  found  in  a  single  stomach  (pi.  22,  fig.  3).  Less 
than  one  per  cent  of  the  grain  found  in  the  stomachs  has  been 
sprouted  grain.  Practically  no  grain  is  found  in  the  stomachs 
during  the  months  of  March  and  July.  In  that  grain  is  more 
slowly  digested  than  insects,  a  smaller  volume  of  grain  is  prob- 
ably consumed  daily. 

Economic  importance. — For  its  destruction  of  sprouting  grain, 
the  western  meadowlark  justly  deserves  criticism.  In  small  fields, 
where  the  birds  are  numerous,  losses  are  great.  The  facts  which 
tend  to  minimize  the  damage  done  are  as  follows:  Much  of 
the  grain  found  in  the  stomachs  is  wild  oats.  Since  cultivated 
oats  always  contains  more  or  less  wild  oats  in  California,  it  can- 
not be  said  that  injury  does  not  result  from  the  destruction 
of  the  latter.  However,  much  of  that  eaten  must  be  taken  in 
places  where  no  injury  results.  All  of  the  grain  taken  in  the 
months  of  August,  September,  and  October  must  be  considered 


1914]   Bryant:  Economic  Status  of  the  Western  Meadoiclark     423 

waste  grain  and  of  little  economic  importance.  Meadowlarks  are 
not  known  to  attack  heads  of  grain.  Whatever  field  grain  is 
taken  is  picked  up  from  the  ground.  Damage  done  to  sprouting 
grain  can  result  only  during  a  limited  period  of  time  (two  weeks). 
After  the  second  leaf  appears  no  damage  can  result. 

Weed  Seed 

Weed  seed  evidently  forms  the  principal  part  of  the  vegetable 
diet  of  this  bird  where  or  when  grain  is  not  available.  The  seeds 
of  such  weed  pests  as  tarweed,  mustard,  tumbleweed,  Napa  thistle, 
pigweed,  amaranth,  canary  grass,  Johnson  grass,  foxtail,  and 
sunflower  are  consumed  in  large  quantities.  The  seeds  of  such 
forage  plants  as  the  burr-clover  and  filaree  are  commonly  eaten. 
The  seeds  of  filaree  (Erodium  cicutarium)  form  the  largest  per- 
centage of  the  weed  seeds  taken  as  food. 

A  stomach  has  seldom  been  found  completely  filled  with  weed 
seed,  for  some  sort  of  grain,  especially  wild  oats,  is  nearly  always 
available  with  the  weed  seeds.  Nevertheless  during  the  late  fall 
weed  seeds  make  up  a  considerable  part  (twenty  per  cent)  of 
the  diet. 

Grass  has  been  occasionally  found  in  the  stomachs.  Whether 
or  not  it  was  taken  intentionally  it  is  impossible  to  state.  Small 
sprouts  from  sprouting  grain  and  sprouting  seeds  have  been 
found  in  some  instances.  As  a  rule  the  seeds  appear  to  be  sep- 
arated from  the  large  sprouts  when  eaten.  Small  pieces  of  straw 
and  other  vegetable  fiber  found  in  the  stomachs  can  be  classified 
as  rubbish  picked  up  with  the  food. 

There  has  been  a  slight  complaint  that  meadowlarks  damage 
sugar  beets  by  feeding  on  the  sprouting  seeds.  Mr.  F.  J.  McCoy, 
assistant  manager  of  the  Union  Sugar  Company,  Betteravia, 
California,  says  on  this  point:  "I  have  noticed  meadowlarks 
in  early  spring  in  our  beet  fields,  but  noticed  they  were  feeding 
on  insects. ' '  Stomach  examination  has  failed  to  disclose  any  beet 
seeds.  The  stomachs  of  birds  collected  in  beet  fields  have  been 
found  filled  with  insects. 

Quantity  destroyed. — The  maximum  consumption  *  of  weed 
seed  occurs  in  October,  when  nearly  one-fourth  of  the  food  is 
made  up  of  this  kind  of  food.  Weed  seed  amounts  to  five  and 


424          University  of  California  Publications  in  Zoology    ["VOL.  11 

three-tenths  per  cent  of  the  food  for  the  year.  Over  150  seeds 
of  filaree  (Erodium  sp.)  have  been  taken  from  a  single  stomach. 
Tarweed,  pigweed,  tumbleweed,  mustard,  turkey  mullein,  Napa 
thistle,  Johnson  grass,  canary  grass,  foxtail,  sunflower,  burr- 
clover,  and  nightshade  seeds  have  been  found  in  numbers  ranging 
up  to  fifty.  In  some  few  instances  stomachs  have  been  found 
entirely  filled  with  weed  seeds.  Western  meadowlarks  appear 
to  feed  upon  weed  seeds  to  a  considerable  extent  during  the  time 
in  which  they  are  available.  Most  weed  seeds  do  not  mature 
until  late  summer  and  fall.  After  plowing  begins  they  are  no 
longer  available  in  cultivated  districts,  except  along  fence  rows 
and  in  uncultivated  fields. 

Economic  importance. — The  destruction  of  weed  seeds  must 
be  considered  of  value  to  the  agriculturist.  Weeds  even  in  small 
numbers  take  a  toll  in  the  grain  field.  The  destruction  of  weed 
seeds  accomplished  by  western  meadowlarks  must  help  to  limit, 
in  some  measure,  the  number  of  weeds  which  grow  in  fields  and 
fence  rows  the  following  year.  Meadowlarks  feeding  in  grain 
fields  must  destroy  weed  seed  that  would  not  otherwise  be  de- 
stroyed. Their  habit  of  feeding  on  sprouting  seeds  increases 
their  efficiency  as  weed-seed  destroyers.  Seeds  eaten  are  digested. 
In  no  case  have  undigested  seeds  been  found  in  excrement. 

Fruit 

No  vegetable  matter  found  in  the  stomachs  has  been  identified 
as  fruit.  Grape  seeds  have  been  found  in  a  number  of  cases  and 
there  is  no  doubt  that  western  meadowlarks  eat  grapes  to  a  slight 
extent.  No  serious  complaint  as  to  their  depredations  in  this 
direction  has  been  received.  The  stomachs  of  practically  all  of 
the  birds  collected  in  vineyards  have  been  filled  with  insects, 
mostly  beetles. 

A  systematic  list  of  the  grain  and  weed  seeds  found  in  the 
stomachs  follows : 

Grain 
Oats 
Barley 
Wheat 

Field  corn  (Zea  mays) 
Sorghum  (Andropogon  sorghum  subsp.) 


1914]   Bryant:  Economic  Status  of  the  Western  Meadowlark     425 

Fruit 

Grape  seeds  (Vitis  sp.) 

Weed  seed 

Gramineae  (Grass  family) 

Andropogon  sorghum  halepensis.    Johnson  grass 

Panicum  sp.     Panic  grass 

Chaetochloa  glauca.     Bristly  foxtail 

Chaetochloa  viridis.     Foxtail 

Phalaris  minor.     Small  canary  grass 

Avena  fatua.    Wild  oats 

Bromus  sp.     Brome  grass 

Lolium  temulentum.     Darnel 

Hordeum  sp.     Barley  grass 
Cyperaceae  (Sedge  family) 

Carex  sp.     Sedge 
Polygonaceae  (Buckwheat  family) 

Rumex  sp.     Dock 

Polygonum  sp.     Knotweed 
Chenopodiaceae  (Goosefoot  family) 

Chenopodium  sp.     Goosefoot  pigweed 
Amarantaceae  (Amaranth  family) 

Amaranthus  graecizans.     Tumbleweed 

Amaranthus  sp.     Amaranth 
Portulacaceae  (Purslane  family) 

Lewisia  sp.     Bitter  root 
Eanunculaceae  (Buttercup  family) 

Ranunculus  sp.     Buttercup 
Cruciferae  (Mustard  family) 

Brassica  nigra.    Black  mustard 

Brassica  sp.     Mustard 
Leguminosae  (Pea  family) 

Medicago  hispida.     Burr  clover 

Medicago  arabica.     Spotted  medick 

Melilotus  indica.     Yellow  melilot 

Melilotus  sp.     Sweet  clover 

Trifolium  sp.     Clover 

Lupinus  sp.     Lupine 
Geraniaceae  (Geranium  family) 

Erodium  cicutarium.     Red-stem  filaree 

Erodium  sp.     Filaree 
Euphorbiaceae  (Spurge  family) 

Eremocarpus  setigerus.     Turkey  mullein 
Malvaceae  (Mallow  family) 

Sida  hederacea.     Alkali  mallow 
Onagraceae  (Evening  Primrose  family) 

Oenothera  ovata.     Golden  eggs 
Primulaceae  (Primrose  family) 

Anagallis  arvensis.     Pimpernel 


426  University  of  California  Publications  in  Zoology    [VOL.  11 

Boraginaceae  (Borage  family) 

Amsinckia  intermedia.     Amsinckia 

Amsinckia  sp.     Amsinckia 

Cynoglossum  sp.    Hound's  tongue 
Eubiaceae  (Madder  family) 

Galium  sp.     Bedstraw 
Solanaceae  (Nightshade  family) 

Solanum  sp.     Nightshade 
Compositae  (Sunflower  family) 

Lactuca  scariola.    Prickly  lettuce 

Centaurea  melitensis.     Napa  thistle 

Centaurea  solstitialis.     Barnaby's  thistle 

Carduus  sp.     Thistle 

Hemizonia  sp.     Tarweed 

Helianthus  annuus.     Common  sunflower 

Iva  axillaris.     Kagweed 

Economic  importance  of  vegetable  food. — The  destruction  of 
sprouting  grain  means  a  loss  of  dollars  and  cents  to  the  rancher. 
This  loss  is  minimized  somewhat  by  the  limited  time  during 
which  injury  is  possible  and  the  possibility  of  protective  measures. 
A  much  smaller  loss  can  be  attributed  to  the  destruction  of  grain 
picked  up  in  newly  sown  fields.  Grain  on  or  near  the  surface 
of  the  ground  in  seeded  fields  is  of  doubtful  value,  as  it  cannot 
be  depended  upon  to  furnish  a  strong,  healthy  plant.  Much  of 
the  wild  oats  and  some  of  the  tame  oats  must  be  considered  waste 
grain  or  uncultivated  grain.  Its  destruction,  in  spite  of  its 
utility  as  feed,  cannot  be  considered  a  direct  loss  in  money  value. 

Practically  all  the  other  seeds  destroyed  are  the  seeds  of  weed 
pests.  The  destruction  of  the  seeds  of  certain  forage  plants  such 
as  filaree  could  be  considered  a  detriment  if  they  were  destroyed 
in  large  enough  quantities  to  make  any  difference  in  the  amount 
of  forage  available.  This  same  plant  is  considered  a  weed  in 
many  places.  Any  destruction  of  the  seeds  of  thistles,  sunflowers, 
Johnson  grass  and  like  weeds  must  be  considered  a  benefit. 


ANIMAL  FOOD 
Coleoptera  (Beetles) 

The  most  constant  article  of  diet  of  the  meadowlark  consists 
of  beetles.  The  habitat  of  the  bird  would  forecast  this  fact. 
Ground-beetles  (Carabidae,  Tenebrionidae) ,  click-beetles  (Elate- 


1914]   Bryant:  Economic  Status  of  the  WesternMeadoivlark     427 


ridae),  and  weevils  (Rhynchitidae,  Calandridae,  Otiorhynchidae) 
form  the  largest  per  cent  of  this  kind  of  food.  Representatives 
of  practically  every  family  of  the  Coleoptera,  however,  have  been 
found  in  the  stomachs. 

A  svstematic  list  of  the  beetles  identified  follows : 


Cincindelidae 

Cincindela  sp. 
Carabidae 

Calosoma  sp. 

Amara  californica  Dej. 

Amara  conflata  Lee. 

Calathus  ruficollis  Dej. 

Platinus  bicolor  Lee.  (?) 
Staphylinidae 

Staphylinus  tarsalis  Mann. 

Creophilus  villosus 
Silphidae 

Silpha  ramosa  Say 
Dermestidae 

Dermestes  sp. 
Erotylidae 

Languria  sp. 
Histeridae 

Saprinus  fimbriatus  Lee. 
Buprestidae 


Elateridae 

Cardiophorus  tenebrosus  Lee. 

Anchastus  einereipennis  Mann. 

Drasterius  livans  Lee. 

Drasterius  sp. 

Megapenthes  aterrimus  Horn 

Dolopius  lateralis 

Melanotus  variolatus  Lee. 

Limonius  infuscatus  Mots. 

Limonius  canus  Lee. 

Limonius  californicus  Mann. 
Lampyridae 

Telephorus  censors  Lee. 
Malachidae 

Collops  marginellus  Lee. 
Scarabaeidae 

Aphodius  subaeneus  Lee. 

Aphodius  granarius  Linn. 

Aphodius  rugifrons  Horn. 

Hoplia  sp. 

Cotalpa  ursina  Horn 

Ligyrus  gibbosus  De  Greer 


Harpalus  pennsylvanicus  Dej. 

Agonostoreus  maculatus  Lee. 

Pterostichus  sp. 

Bladyeellus  rupestris  Say 

Anisodactylus  dilatatus  Dej. 

Anisodactylus  sp. 
Dytiscidae 

Agabus  lugens  Lee. 
Chrysomelidae 

Glyptoscelis  albidus  Lee.  (?) 

Gastroidea  sp. 

Diabrotica  soror  Lee. 

Disonycha  sp. 

Chaetocnema  sp. 

Microrhopala  melsheimeri  Cr. 
Tenebrionidae 

Blapstinus  gregalus  Casey 

Blapstinus  rufipes  Casey 

Blapstinus  elongatus  Casey 

Coniontis  subpubescens  Lee. 

Coniontis  viatica  Esch. 

Eulabis  pubescens  Lee. 

Eurymetopon  sp. 

Eleodes  sp. 
Meloidae 


Otiorhynchidae 

hnigopsis  effracta  Lee. 
Rhigopsis  sp. 


Curculionidae 

Sitones  californicus  Fah. 

Sitones  sordidus  Lee. 

Lixus  perferatus  Lee. 

Cleonus  virgatus  Lee. 

Centrocleonus  near  angularis  Lee. 

Baris  cuneipennis  Casey 

Baris  sp. 
Calandridae 

Sphenophorus  vomerinus  Lee. 

Sphenophorus  simplex  Lee. 

Sphenophorus  sp. 


428  University  of  California  Publications  in  Zoology    [VOL.  11 

Quantity  destroyed. — Beetles  are  taken  every  month  of  the 
year,  and  form  21.3  per  cent  of  the  total  food.  Stomachs  have 
often  been  found  filled  with  nothing  but  beetles.  From  twenty 
to  fifty  have  been  found  in  a  single  stomach. 

Wireworms,  the  larvae  of  click-beetles  (Elateridae),  are  taken 
in  large  numbers  where  they  are  available.  As  they  are  less 
often  seen  above  ground  than  cutworms,  it  is  only  natural  that 
they  do  not  form  nearly  so  large  a  percentage  of  the  food  for 
the  year.  The  adult  click-beetles  are  also  taken  (see  pi.  24,  fig.  8) . 

Economic  importance. — Wireworms  are  injurious  to  the  roots 
of  plants.  Damage  by  them  to  garden  truck  and  pasture  land 
is  of  common  occurrence.  The  ability  of  the  meadowlark  to 
probe  into  and  remove  from  the  soil  such  insects  increases  its 
value  as  an  insect  destroyer.  The  destruction  of  wireworms  must 
be  considered  a  benefit  of  considerable  importance,  especially  in 
meadow  and  pasture  land. 

By  far  the  greater  number  of  beetles  taken  as  food  are  the 
common  ground  beetles  (Carabidae).  These  beetles  are  often 
classified  as  beneficial  insects,  because  they  are  supposed  to  feed 
on  other  injurious  insects.  Certain  ones  are  predacious  and  are 
known  to  feed  on  fly  and  beetle  larvae  in  California.  Of  the  food 
habits  of  others  little  is  definitely  known,  and  we  are  justified 
in  speaking  of  them  as  neutral,  for  they  do  practically  no  harm 
and  are  not  known  to  do  any  particular  good.  Tiger-beetles 
(Cincindelidae)  and  carrion-beetles  (Staphylinidae),  eaten  to 
some  extent,  must  be  numbered  among  the  beneficial  beetles  de- 
stroyed. The  meadowlark,  however,  does  feed  upon  many  injur- 
ious beetles,  chief  of  which  are  click-beetles  (Elateridae),  pina- 
cate  beetles  (Eleodes  sp.),  leaf -beetles  (Chrysomelidae),  snont- 
beetles  (Otiorhynchidae,  Curculionidae),  and  weevils  (Calan- 
dridae).  Among  the  leaf-beetles  is  numbered  the  destructive 
California  flower-beetle  (Diabrotica  soror).  This  and  other 
members  of  the  family  constitute  some  of  the  worst  beetle  enemies 
of  our  crops.  Snout-beetles  (curculios)  and  weevils  (Spheno- 
phorus  sp.)  are  well-known  pests  of  fruit  and  grain.  The  con- 
tinual destruction  of  large  numbers  of  these  injurious  beetles 
must  be  considered  a  decided  benefit. 


1914]   Bryant:  Economic  Status  of  the  Western  Meadowlark     429 

Orthoptera  (Grasshoppers  and  Crickets) 

Grasshoppers  and  crickets  form  a  large  percentage  of  the 
meadowlark's  food  during  the  summer  and  fall,  making  up  as 
high  as  eighty-five  per  cent  of  the  food  in  August.  The  species 
of  grasshopper  taken  is  the  one  most  available.  Practically  all 
of  the  species  found  in  the  state  are  doubtless  represented  in  the 
stomachs.  The  common  cricket  (Gryllus)  and  the  Jerusalem 
cricket  (Stenopelmatus  sp.)  are  common  articles  of  diet.  Katy- 
dids (Locustidae),  being  less  common  insects  and  having  a 
different  habitat,  are  not  taken  so  often. 

A  systematic  list  of  the  Orthoptera  found  in  the  stomachs 
follows : 

Gryllidae  Arphia  sp. 

Gryllus  integer  Scud.  Dissosteira  spurcata  Saus. 

Gryllus  pennsylvanicus  Burm.  Conozoa  behrensi  Saus. 

Locustidae  Aeridiinae 

( ?)  Macrocentrum  sp.  Oedaleonotus  enigma  Scud. 

Conocephalus  acutalus  Scud.  Melanoplus  devastator  Scud. 

Stenopelmatus  irregularis  Brun.  Melanoplus  differentialis  Uhler 

Stenopelmatus  sp.  (  ?)  Melanoplus  unif ormis  Scud. 

Acridiidae  Fulgoridae 

Cedipodinae  (?)  Labia  minor 

Camnula  pellucida  Scud. 


Quantity  destroyed. — Next  to  beetles,  grasshoppers  form  the 
most  important  article  of  diet.  Nearly  fifteen  per  cent  of  the 
food  for  the  year  is  made  up  of  these  insects.  Parts  of  as  many 
as  twenty-six  large  grasshoppers  (one  inch  or  over  in  length) 
and  fifty-eight  small  grasshoppers  (one-half  inch  in  length)  have 
been  found  in  a  single  stomach.  The  maximum  amount  of  this 
food  is  taken  in  June,  July,  and  August,  the  birds  feeding  almost 
exclusively  on  these  insects  during  these  months.  The  state  over, 
some  grasshoppers  are  taken  by  the  meadowlarks  every  month 
in  the  year.  The  quantity  taken  closely  parallels  the  abundance 
of  these  insects. 

Crickets  as  well  as  grasshoppers  are  relished  by  the  meadow- 
lark.  As  many  as  fifteen  pairs  of  mandibles  have  been  taken 
from  a  single  stomach,  showing  that  at  least  fifteen  common  black 


430          University  of  California  Publications  in  Zoology    [  VOL.  11 

crickets  (Gryllus  pennsylvanicus)  had  been  eaten  within  four 
hours  by  that  particular  bird.  Five  per  cent  of  the  food  for  the 
year  is  made  up  of  crickets.  Wood  crickets,  better  known  as 
Jerusalem  crickets  (Stenopelmatus  sp.),  being  less  abundant  than 
the  common  cricket,  are  less  often  taken  as  food. 

Economic  importance. — Grasshoppers  can  be  classed  as  injur- 
ious insects.  The  extent  of  their  damage  can  be  traced  to  their 
abundance  rather  than  to  the  presence  of  any  particular  kind 
of  grasshopper.  The  species  which  most  often  become  abundant 
enough  to  cause  serious  losses  in  this  state  are  the  differential 
grasshopper  (Melanoplus  differentialis) ,  the  pale-winged  grass- 
hopper (Melanoplus  uniformis),  the  devastating  grasshopper 
(Melanoplus  devastator),  and  the  valley  grasshopper  (Oedaleo- 
notus  enigma).  All  of  these  grasshoppers  are  destroyed  in  great 
numbers  by  the  meadowlark.  The  more  abundant  these  insects 
become,  the  more  do  these  birds  turn  their  attention  to  this  kind 
of  food.  Where  grasshoppers  are  abundant,  meadowlarks  have 
been  found  to  average  as  high  as  fifty  grasshoppers  a  day.  (See 
Bryant,  1912d.)  As  a  grasshopper  destroyer,  the  meadowlark  is 
unequaled  by  any  other  bird  unless  it  be  the  blackbird,  and  then 
only  because  of  greater  numbers  of  blackbirds.  As  grasshopper 
outbreaks  continue  to  ravage  certain  parts  of  the  state  each  year, 
the  meadowlark  performs  a  service  to  agriculture  that  can  hardly 
be  overestimated,  in  that  it  helps  to  keep  the  insects  down  to 
normal  numbers,  so  that  losses  do  not  result,  and  prevents  greater 
losses  by  taking  a  greater  toll  at  the  time  of  an  outbreak. 

Crickets  are  usually  classed  as  injurious  insects.  The  degree 
of  injury,  as  with  the  grasshoppers,  depends  largely  upon  their 
abundance.  Since  the  species  of  crickets  fed  upon  by  the  mead- 
owlark feed  almost  entirely  upon  plants  and  are  often  destructive 
to  grain,  their  destruction  is  to  be  desired  by  the  rancher.  This 
is  especially  the  case  with  the  Jerusalem  cricket,  which  is  very 
destructive  to  potatoes. 

Lepidoptera  (Butterflies  and  Moths) 

The  general  law  that  birds  do  not  eat  butterflies  to  any  great 
extent  appears  to  hold  good  in  the  case  of  the  western  meadow- 
lark.  (See  p.  481).  However,  the  following  dependable  obser- 


1914]   Bryant:  Economic  Status  of  the  Western  Meadowlark     431 

vation  made  by  Mr.  John  G.  Tyler  of  Fresno,  California,  also 
furnishes  evidence  of  the  fact  that  butterflies  are  occasionally, 
at  least,  destroyed.  "While  strolling  along  the  road  east  of  this 
city  the  writer  noticed  a  field  of  alfalfa  that  was  infested  with 
yellow  butterflies.  A  nearer  approach  revealed  the  presence  of 
several  meadowlarks,  and  I  was  so  fortunate  as  to  see  one  of 
these  birds  seize  a  butterfly  and  make  way  with  it.  I  am  not 
prepared  to  say  that  the  victim  was  actually  swallowed,  but  it 
was  certainly  captured  and  killed." 

The  larvae  of  butterflies  and  moths  are  common  articles  of 
diet.  Cutworms  and  caterpillars  form  ten  per  cent  of  the  food 
for  the  year,  reaching  a  maximum  in  May  and  June,  when  they 
amount  to  nearly  a  third  of  the  meadowlark's  food. 

Even  hairy  caterpillars  do  not  escape  destruction.  In  one 
instance  the  larva  of  the  mourning-cloak  butterfly  (Euvanessa 
antiopa)  has  been  found  in  the  stomach.  Smaller  hairy  cater- 
pillars are  of  common  occurrence.  Both  the  larva  and  pupa  of 
the  sphinx  moth  have  been  taken  from  stomachs.  Pupae  do  not 
form  so  important  a  part  of  the  diet  as  do  the  larvae. 

The  only  Lepidoptera  positively  identified  follow: 


LEPIDOPTERA  (BUTTERFLIES  AND  MOTHS) 

Noctuidae  Nymphalidae 

Peridromia  sp.  Eugonia  calif ornica  (Boisd.) 

(?)  Euvanessa  antiopa  (Linn.) 

-(?)  (?)  Papilio  sp. 

Sphingidae 

(?)  Phlegethontius  sp. 


Quantity  destroyed. — Cutworms  and  caterpillars  form  about 
twelve  per  cent  of  the  food  for  the  year.  The  maximum  quantity 
is  taken  in  the  months  of  March  and  April,  when  almost  half  of 
the  food  taken  is  made  up  of  these  insects.  Many  of  the  stomachs 
contained  as  many  as  twenty  large  cutworms  or  caterpillars. 
One  bird  collected  at  Red  Bluff,  Tehama  County,  contained  sixty- 
six  cutworms  and  over  thirty  small  beetles  (pi.  23,  fig.  5). 


432          University  of  California  Publications  in  Zoology    [VOL.  11 

Economic  importance. — Cutworms  and  army  worms  can  be 
classed  as  two  very  important  pests  in  California.  Garden  truck 
and  even  trees  are  sometimes  defoliated  when  these  insects  become 
numerous.  The  depredations  of  the  grape  cutworm  are  only  too 
well  known  in  the  state.  Caterpillars  are  vegetable  feeders  and 
are  always  classed  as  injurious.  The  destruction  of  these  pests 
in  very  large  quantities  by  the  western  meadowlark  must  cause 
a  direct  saving  to  the  rancher  and  fruit  grower. 


Hemiptera  (Bugs) 

Stink-bugs  (Pentatomidae)  appear  to  be  relished  in  spite  of 
their  excretions,  for  they  are  taken  in  large  numbers.  Squash- 
bugs  (Anasa  sp.)  have  been  found  in  only  a  few  instances. 
Negro-bugs  (Corimelaena)  form  the  only  other  important  Hem- 
iptera taken.  Cicadas  appear  to  be  relished  and  often  caught. 
Two  stomachs  have  contained  aphids  (Aphis  brassicae).  The 
following  Hemiptera  have  been  identified : 

HEMIPTERA  (Buos) 

Jassidae  Coreidae 

—  ?  Corizus  sp. 

Aphidae  Alydus  pliosulus 

Aphis  brassicae  Linn.  Anasa  sp. 

Membracidae  Pentatomidae 

Stictocephala  franciscana  Stal.  Podisus  pallens  Stal. 

Cicadidae  Podisus  sp. 

Platypedia  areolata  Uhl.  Eusehistus  conspersus  Uhl. 

Platypedia  minor  Uhl.  Eusehistus  servus  Say 

Eeduviidae  Corimelaenidae 

—  ?  Corimelaena  sp. 


Quantity  destroyed. — The  commonest  true  bugs  destroyed  by 
western  meadowlarks  are  stink-bugs  (Pentatomidae),  negro-bugs 
(Cormelaenidae),  leafhoppers  (Jassidae),  and  cicada  flies  (Cica- 
didae). They  form  nearly  two  per  cent  of  the  food  for  the  year. 
As  many  as  twenty  stink-bugs  have  been  taken  from  a  single 


1914]    Bryant:  Economic  Status  of  the  Western  Meadowlark     433 

stomach.  Negro-bugs  are  not  taken  in  such  large  numbers,  nor 
are  they  so  abundant.  Two  stomachs  have  been  found  almost 
filled  with  leafhoppers,  and  many  others  contained  two  to  ten 
of  these  insects.  Cicadas,  near  relatives  of  the  eastern  seventeen- 
year  locust,  are  occasionally  taken,  probably  as  often  as  they  are 
available.  Bugs  form  over  three  per  cent  of  the  food  for  the  year. 
Economic  importance. — Most  stink-bugs  are  vegetable  feeders 
and  occasionally  give  trouble.  Negro-bugs  are  troublesome  on 
berries.  In  the  destruction  of  these  insects  the  western  meadow- 
lark  is  also  conferring  a  benefit.  Leafhoppers  are  injurious  to 
plants,  because  they  secure  their  food  by  sucking  the  juice  of 
the  plant.  Any  destruction  of  leafhoppers,  however  small,  is 
of  value.  The  cicada  in  California  is  not  abundant  enough  to 
be  of  economic  importance.  It  lays  its  eggs  in  the  sapwood  of 
plants  and  trees.  Since  these  insects,  if  they  became  abundant, 
would  cause  trouble  as  does  the  seventeen-year  locust  of  the  east, 
their  destruction  is  to  be  looked  upon  with  favor. 


Hymenoptera  (Ants.  Bees,  and  Wasps) 

Ants  appear  to  be  taken  irrespective  of  size  or  kind,  for  they 
are  to  be  found  from  the  smallest  to  the  largest.  The  common  red 
and  black  ants  (Messor,  Pogonomyrmex) ,  field  ants  (Formica), 
and  carpenter  ants  (Camponotus)  are  most  abundant  in  the 
stomachs. 

Ichneumon  flies  are  taken  in  considerable  numbers.  Bees 
and  wasps  form  a  less  percentage  of  the  food  made  up  of  Hymeno- 
ptera. In  but  one  or  two  cases  was  a  bee  definitely  identified 
as  a  honey  bee  (Apis  mellifera).  Solitary  bees  (Chrysis)  and 
even  bumblebees  (Bombus  calif ornicus)  have  been  found.  Cow- 
killers  (Mutillidae)  are  occasionally  eaten,  although  it  has  com- 
monly been  supposed  that  they  were  well  protected  from  attack 
by  their  sting,  hairy  covering,  and  warning  coloration. 

For  several  reasons  the  Hymenoptera  have  been  very  difficult 
to  identify.  The  finely  comminuted  condition  of  the  insects  has 
proved  an  almost  insurmountable  difficulty.  The  following:  only 
have  been  identified: 


434          University  of  California  Publications  in  Zoology    [VOL.  11 

HYMENOPTERA  (ANTS,  BEES,  ANb  WASPS) 
Ichneumonidae  Vespidae 

?  Polistes  aurifer  Sauss. 

Polistes  minor  Beauv. 
-  ?  Sphegidae 

1  Ammophila  sp. 

Mutillidae  Formicidae 

Sphaerothalma  californica  Formacinae 

Sphaerothalma  aureola  Camponotus   sp. 

Apidae  Formica  sp. 

( ?)  Apis  mellif era  Dolichoderinae 

Bombidae  Tapinoma  sessile  Say 

Bombus  californicus  Smith  Myrmicinae 

Chrysididae  Messor  andrei  Mayr. 

Chrysis  sp.  Pogonomyrmex    californicus 

Buckley 
Pogonomyrmex  sp. 

Quantity  destroyed. — Bees  and  wasps  form  3.6  per  cent  of 
the  yearly  food.  In  no  case  have  ichneumon  flies,  which  are 
valuable  parasitic  insects,  been  taken  in  numbers,  five  being  the 
maximum  found  in  a  single  stomach.  Their  rapid  flight  prob- 
ably prevents  a  greater  toll  being  taken.  Ants  are  often  eaten 
in  large  quantities  and  form  over  two  per  cent  of  the  food  for 
the  year.  It  is  not  an  uncommon  occurrence  to  find  a  stomach 
almost  filled  with  ants.  Over  one  hundred  have  been  found  in 
a  single  stomach. 

Economic  importance. — Most  bees  and  wasps  are  considered 
beneficial  insects.  Ants  are  either  injurious  or  neutral;  few  are 
beneficial.  In  the  destruction  of  ichneumon  flies  the  western 
meadowlark  is  destroying  a  valuable  parasitic  insect.  The  de- 
struction of  bees  and  wasps  must  also  be  reckoned  as  a  count 
against  the  bird.  However,  the  small  numbers  destroyed  mini- 
mize greatly  the  real  and  the  possible  damage  done.  The 
destruction  of  most  kinds  of  ants  makes  little  difference  one  way 
or  the  other,  owing  to  their  abundance  and  scavenger  habits. 

Dipt  era  (Flies} 

A  few  members  of  the  family  Muscidae,  a  few  flower-flies 
(Eristalis  sp.)  'and  crane-flies  (Tipula  sp.)  and  the  pupae  of 
syrphid  flies  (Syrphus)  are  the  only  representatives  of  the 
Diptera  which  have  been  found  in  the  stomachs  of  western 
meadowlarks.  The  following  have  been  identified:  • 


1914]   Bryant:  Economic  Status  of  the  Western  Meadowlark     435 

DlPTERA 

Eristalis  tenax  Lucilia  caesar  L. 

Tachina  sp.  Musca  sp. 

Syrphus  sp.  ?  Tipula  sp. 

Quantity  destroyed. — Flies  do  not  constitute  any  important 
percentage  (about  one  per  cent)  of  the  food  for  the  year  and 
when  found  have  been  in  small  numbers.  A  few  green  bottle- 
flies  and  other  members  of  the  same  family  are  eaten,  as  are  also 
flower-flies  and  drone-flies.  Birds  collected  at  El  Toro,  Orange 
County,  during  1911  had  eaten  large  numbers  of  the  pupae  of 
flower-flies  (Syrphus  sp.) .  Crane-flies  (Tipula  sp.)  are  not  taken 
as  often  as  it  would  seem  they  would  be  from  their  abundance. 
Evidence  is  at  hand,  however,  that  western  meadowlarks  feed 
largely  on  the  larvae  when  they  become  abundant.  Mr.  W.  M. 
Hughes  of  Madera,  Madera  County,  has  made  the  following 
report:  "When  I  visited  the  tract  of  land  affected,  I  found 
myriads  of  blackbirds  and  thousands  of  meadowlarks  on  the 
ground  making  small  holes  into  the  ground  at  the  roots  of  the 
plants  and  taking  out  the  worm.  Several  hundred  acres  of  fine 
crop  was  destroyed  before  the  birds  collected  in  numbers  suffi- 
cient to  destroy  the  pest. ' '  The  outbreak  referred  to  was  in  the 
vicinity  of  Minturn,  Madera  County,  in  1909.  Several  hundred 
acres  of  barley  were  destroyed  by  crane-fly  larvae  at  this  time. 

Economic  importance. — In  spite  of  the  fact  that  the  larvae 
feed  upon  decaying  matter,  most  flies  are  considered  pests  be- 
cause some  of  them  carry  germs  of  disease.  Green  bottle-flies 
are  disease  carriers.  The  larvae  of  flower-flies  feed  upon  plant 
lice  and  hence  are  considered  beneficial.  The  larvae  of  crane- 
flies,  on  the  other  hand,  are  destructive  to  vegetation.  The  small 
numbers  of  flies  taken  and  the  fact  that  injurious  as  well  as 
beneficial  forms  are  eaten  make  the  destruction  of  Diptera  by 
meadowlarks  of  little  consequence. 

Arachnida  (Spiders) 

Quantity  destroyed. — Spiders  and  their  egg-cases  form  less 
than  one  per  cent  of  the  food  for  the  year.  Most  of  the  spiders 
taken  are  grass  spiders  (Agalenidae)  and  daddy-long-legs  (Pha- 


436  University  of  California  Publications  in  Zoology    [VOL.  11 

langidae),  the  commonest  of  those  found  in  pastures.  In  no 
instance  have  more  than  two  spiders  been  taken  from  the  same 
stomach. 

Economic  importance. — Spiders  should  be  considered  as  doubt- 
fully beneficial  or  of  neutral  value  to  the  agriculturist,  in  spite 
of  the  fact  that  they  feed  largely  on  insects.  As  a  rule  spiders 
are  not  abundant  enough  to  be  of  great  economic  importance. 
Their  destruction  at  the  hands  of  the  meadowlark  is  of  no  conse- 
quence, as  the  resulting  effect  on  insect  life  is  so  small. 

Miscellaneous  Animal  Food 

Miscellaneous  articles  of  diet  form  three  and  one-half  per 
cent  of  the  food  for  the  year.  The  common  sow-bug  (Porcellio 
scaber)  is  the  commonest  crustacean  found  in  stomachs.  Two 
birds  had  eaten  snails.  Two  birds  from  San  Diego  had  each 
taken  a  scorpion.  But  few  earthworms  have  been  found  in  the 
stomachs.  Centipedes  (Scolopendra  sp.)  and  millipedes  (Julus) 
appear  to  form  a  constant  part  of  the  diet.  They  are  evidently 
taken  regularly  where  available.  Two  birds  had  eaten  ant-lions 
(Myrmeleon  sp.). 

Quantity  destroyed. — Centipedes,  millipedes,  scorpions,  ant- 
lions,  and  sow-bugs  may  be  considered  oddities  in  the  diet.  Their 
slight  availability  may  account  in  some  measure  for  the  small 
numbers  taken  by  the  western  meadowlark. 

Economic  importance. — Of  these  miscellaneous  elements  in  the 
diet,  only  millipedes  and  sow-bugs  can  be  considered  injurious. 
Centipedes  are  usually  considered  beneficial,  scorpions  injurious, 
and  ant-lions  of  neutral  value.  None  of  these  forms  is  taken 
in  sufficient  quantity  to  make  their  destruction  either  an  injury 
or  a  benefit. 

Inorganic  Matter 

Pebbles,  used  for  grinding  the  food,  make  up  the  inorganic 
matter  found  in  the  stomachs.  White  and  red  pebbles,  probably 
because  of  their  conspicuousness,  predominate.  Pebbles  appear 
to  be  necessary  as  an  aid  to  the  digestion  of  grain,  but  much  less 
necessary  for  the  digestion  of  insects.  Pebbles  are  nearly  always 


1914]   Bryant:  Economic  Status  of  the  WesternMeadowlark     437 

found  with  grain  and  weed  seed,  but  seldom  with  insect  food. 
Doubtless  the  chitinous  parts  of  the  insects  largely  take  their 
place. 

Two  nestlings  and  two  adults  contained  parts  of  egg-shells. 
It  is  a  well-known  fact  that  birds  often  eat  the  broken  shells 
after  the  young  have  hatched.  In  one  instance,  at  least,  the 
parent  birds  had  fed  the  young  on  the  shells. 

PRINCIPAL,  ARTICLES  OF  DIET 

The  kinds  of  food  forming  a  definite  part  of  the  food  of  the 
western  meadowlark  for  the  year  are  as  follows : 

VEGETABLE  ANIMAL 

Grain                            30.8%                     Coleoptera  21.3% 

Weed  seed                     5.3                         Orthoptera  20.3 

Miscellaneous                 .6                         Lepidoptera  12.2 

Hemiptera  1.7 

Hymenoptera  5.6 

Diptera  .1 

Arachnida  .2 
Miscellaneous  insects   1.9 

EXAMINATION  OP  FECES 

Other  than  stomach  examination,  the  examination  of  feces 
would  appear  to  give  the  best  evidence  as  to  the  food  of  birds. 
That  a  considerable  amount  of  knowledge  concerning  the  food 
of  meadowlarks  can  be  obtained  in  this  way  is  evidenced  by  the 
fact  that  the  examination  of  feces  of  meadowlarks  collected  June 
15,  1912,  in  the  Berkeley  Hills  showed  the  following:  Thorax 
of  spider,  heads  of  ants,  mandibles  and  other  parts  of  grass- 
hoppers, wing  covers  and  mandibles  of  beetles  and  pubescence 
from  wild  oats. 

The  examination  of  some  feces  collected  from  nestling  birds 
at  Lathrop,  San  Joaquin  County,  showed  that  these  same  hard 
parts  passed  through  the  digestive  tract  undigested.  Conse- 
quently this  affords  a  practical  method  of  determining  the  kind 
of  food  taken.  Its  value  as  a  means  of  determining  the  amount 
of  food  is  much  less,  for  only  the  more  resistant  parts  can  be 
found. 


438          University  of  California  Publications  in  Zoology    [VOL. 11 


pOOJ        TH     TH     L^     rH     O     b-    OS  O     rH     GO     rH     GO     b-     CM     CM     IO     CM    CO 

tr.ir>T      rH     O*     O     rH*     IO"     O     CO*  1O°     i-H     OS*     OS*     CO     CM*     OS*     TH     CD     TH     TH 

COCOTHlOCOlOCOOSGOlOCMTHrH  IOTHCO 

CO     CO    CO    OS     O    CO     rH  O    OS     CM     OS     CM     CO     CO     00     IO     b--     b- 

oo'  os*  os*  06  10'  os"  co  10"  ao*  o°  o  co*  t>*  o  10*  co  10'  10* 

COCOIOTHCOTHCO  rHTHb-lOOOOSTHlOCDOS 

CM  oo  co  co   :   :  os  :    co  TH  co  co  b-;  os  CM  oo  co 

rH     CM*        '     rH         j         !     TH*  j              CM*                      rH     rH     CM*                      CO 

(epiuq0B.iy)      oq  CM   TH      :   CM   CM   CM  :            :   CM   I-H   o      :   iq   r-i   I-H   iq 

sjgpidg                                      rH  :            : 

iq  os   co   CM   co  TH   b-;  :          TH   oo   TH   os   iq   to   TH   o   TH 

*  CO'  j      rH*         CO*  TH*       "  CO*  IO* 

.      b-  O5_  O   1  IO  CM  b^  :     CM  OS  CM  CO  IO  b-  TH  CO  CO 

'   CM*  CO*  1O*    i  rH  TH*  TH  i      rH  rH  rH      TH  CM*         CO* 

pUB  S99^              •  \f£ 

CM  TH  iq   :  iq  co  O  :  rH  co  rH   :  I-H  iq  co  co  co  b- 

"cOCMICMb-CM*  rH         rH      lo'rHrH 

IO  CM  CO  TH  TH  TH  TH  !  CD  CO  OS  b-  TH  IO  rH  b-  b-  OS 

IO*  CO  CO  IO'  IO'  CD  OS*  !  CO  GO'  CO  TH  OS*  b-*  TH  CO*  rH  OS* 

i-H  rH      i— I  rH  CMrHrHrH 

CO 

ICMOSO      :      i      i  iiooo      ;rHio      icoiq      i   c > 

*  rH    !    I    I  j  CM*       !  r-l 

b-  OS_  CO  CM  rH  CO  O  |  O  IO  CO  TH  O   :  IO  iq  TH  CM 

os*  CM*  10*  CM*  oo'  t^  oo'  :   CM'  TH   o   CM'  co'      :   t-*  b-*  CM*  10" 

rHrHCMCMCM  CM             rH                     CMTHCM 

oocooTHcocooo  :      :oscorHrH      •t>;l^*>;r~J 

",_*,_"           *co*  :CM     "0606      ICM'IO'IO'O 

rH  rH                rH 

O5  TH  CM  CO  TH  rH  O  CO  IO  CO  O  O  TH  IO  b-  IO  OS  CM 

CO*     00*     CO*     b-*     00*     b-"     CM*  IO'     b-^     O     OS*     TH     rH*     rH*     TH     GO     OS     IO' 

COCMCM                                CM  rHCMCMCMCMrH                      CO 

b-COOOSCOrHCM  JrHOSTHCOb-t^lOOOOrH 

"     T-"     TH     TH     O     OS*     ?O  i     b-*     b-*    CO*     CM     CM*     GO     rH     b-^    CM' 

iq  TH  b-  oq  iq  co_  oo  o  os  oo  CM  TH  co   :  cs  TH  co  os 

o  oo*  cd  10*  10*  co  10*  10*  co  co  10  TH  os*   :  r-"  co*  CM*  co 

COCMCOTHCMTHCMOSb-lOrHTH  'THCOCO 

Sq^UOUI        OOOb-CO(MrHb-rHCMGOCOb-cC>'-<OSCMrHCO 

9qUXn^J     I— I                                 rH     rH  rH     rH 

Spaiq        OSOCMOOSCOCMCOOOOSTHOrHCDTHlOCMTH 

™™^     CDCOL-CMIOCOTH  rHOJCMCOCO                      l-b-OS 

rH 

ososososososososososasososososososos 


3  5  -g 

s    «    cr 

CD      CU 


Mco^fy2H02a2rHr^^ia2^^o<1a2 


CO       Oi 

c3     a> 


1914]   Bryant:  Economic  Status  of  the  Western  Meadowlark     439 


POOI 

t^ 

o 

CM 

CM     00 

CM     O 

rH     >0 

r—  i 

,— 

<M    00 

o 

fc» 

9[qB}8S8A  [Bjoj, 

05 

rH 

05 
CO 

O     TH 
»0    10 

CM    CO 

i 

CM 

CO 

QC 
00 

o  co 

CO    TH 

*" 

cd 

CO 

^ 

pooj 

CO 

0 

oo 

00    CM 

00    O 

05    »0 

05 

05 

00    CM 

0 

CO 

,2 

CJ 

IBiniUB  [BJOJ, 

0 
05 

§ 

o 

CO 

05     GO 

£!  S 

05    05 
O5    CO 

CO 

CO 

O5    CO 
CD    LO 

CO 
O5 

CO 

co 

1 

^•p 

CM 

CO 

t- 

rH    CO 

IO    O5 

00    CO 

LO 

IO    t^ 

O 

CD 

i 

Sn08UBU8DSIH 

CO 

oo 

CD 

TH    CO 

IO    CM 

IO    CD 

CM 

CM     TH 

LO 

CM' 

j 

(BptuqoBjy) 

, 

,_, 

CO    IO 

oo  o 

;          ; 

CM 

,_, 

CM    CM 

CM 

sjapidg 

i 

:     : 

! 

rH 

o 

CD 

^    CO 

HH     CM 

00    CM 

fc. 

^ 

10    CM 

0 

% 

s^uy 

LO 

LO 

rH 

CM 

IO 

LO 

rH 

CM' 

to 

ftj 

(  B  ja;aou8uiA  H  ) 

rH 

0 

CM 

co  co 

CO    O 

CO    CM 

GO 

t^ 

O5    CO 

10 

CO 

H 

puB  saag 

1-1 

00 

1-1 

rH     rH 

CO     rH 

rH 

CO 

r~i 

TH 

co' 

> 

(Baa^diuteH) 

^ 

IO    IO 

CO    O5 

10     : 

rH 

t. 

OO    TH 

CO 

^ 

w 

sSng 

i 

rH 

rH     0 

CO    rH 

CO 

r—t 

TH      TH 

rH 

1 

** 

SJBTItdjaiBO 

CD 

•* 

IO    O5 

Tf<    CO 

CO     rH 

Th 

O 

CO    OO 

co 

CM 

o 

puB  sraaoA^iiQ 

^ 

rH 

CO    t^ 

TH      ^ 

GO 

T—  | 

CM 

t-    t- 

O5 

LO 

CM' 

rH 

g 

0 

0<fe»njtnaZ*rf 

OO 

,_, 

I     rH 

t-    rH 

!     rH 

05 

LO 

L-    CM 

05 

™  '      +   n    . 

CO 

CM 

•         l£) 

; 

— 

CO    CM 

h 

Ulo[l>allJ.dJ_ 

' 

1 

O 

8 

LO 

0 

l^. 

0    05 

TH    CO 

co      : 

-r 

0 

05    10 

CM 

s 

s  j9ddoqssBJ-£) 

oo 

CD 

t>*     H- 

CO    05 

oo      ; 

i^. 

CO 

rH     t^» 

O 

TH 

z 

rH 

CM 

TH 

CM 

M 

fa 

rH 

CO 

CD 

(M    t^ 

05    05 

:   o 

CM 

TH 

t^    IO 

rH 

CM 

g 

T^^l       .       kj 

CO 

05 

00    t- 

CO 

!    ^ 

T—  1 

CM    TH 

rH 

IO 

i 

CM 

rH 

I 

(Bja^doaioo) 

LO 

CO 

O 

00    O 

CM    O5 

CO    TH 

CD 

1—  I 

t~    CM 

O 

CO 

s 

S8J}33g 

O5 
10 

CO 

O5 
rH 

00     CO 

IO    IO 
Tfi     CM 

CD    00 
CO    CM 

CM 

CM 

CO    O5 

rH 

CO 
rH 

CM 

w 

>o 

E"1 

CD 

CO     t- 

0    0 

:  LO 

LO 

rH 

05     CO 

CO 

&4 

p^da   f^^^V 

^ 

CQ 

TH 

1    CO 

CO 

CO 

<M 

LO 

O 

O] 

CM 

a 

LO 

CO 

»o  oo 

rH     t~ 

rH     O5 

05 

TH 

rH     TH 

rH 

oo 

^ 

UIBJ£) 

CM 

05 

05     L- 

t-    CM 

CO 

CD 

U3 

rH     CO 

00 

o 

h4 

rH 

CM     rH 

CO 

CM 

CO 

CO    TH 

CO 

O 

A 

sqjuora 

TH 

rH 

rH 

O5    t^ 

CM    CM 

rH     CO 

CM 

CM 

CM    (M 

rH 

a 

jo  jBqxnti^ 

rH 

rH 

T—  ! 

r^ 

rH     iH 

0 

g 

§ 

spjiq 

OO 

rH 

CO 

QC 

t-    0 

CD    IO 

05     rH 
rH     l>- 

CO     TH 

0 

rH 

t^ 

T—  1 

O5     00 
CO    CO 

oo   cc 

rH     i— 

1          O 
CM 

1 

TH 

O5 
( 

**$ 

JB8A 

05 

05 

05 

05    05 

O5     O5 

05     05 

05 

05 

O5     O5 

O5     O 

1 

0 

G 

rrt 

•  rH 

is  E 

M 

O 
43 

c 

Z 

Si 

O 

| 

>  s^ 

*G 
1 

Madera 

Fresno 

S 

"S 

W   ^ 

03 

PQ 

ll 

Kern 

Ventura 

§ 

1 

ff 

& 

Riverside 

O 

0)      •  .H 

be  ft 

:~-       Si 

0   W 

Imperial 

IVf  isr»plln  n 

Is 

.Is 

440          University  of  California  Publications  in  Zoology    LV°L-  n 

QUANTITY  OF  FOOD 

Three  methods  may  be  used  in  estimating  the  quantity  of 
food  in  a  bird's  stomach.  First,  the  articles  found  may  be 
counted;  second,  they  may  be  weighed;  or  third,  they  may  be 
estimated  by  volume. 

The  first  method,  although  important  as  giving  an  idea  of 
the  bird's  economic  value  by  showing  the  number  of  injurious 
insects  or  seeds  destroyed,  fails  to  take  into  account  the  difference 
in  size  of  the  different  articles,  and  does  not  show  the  relative 
amounts  of  each  kind.  The  second  method  has  been  generally 
disregarded  because  of  its  impracticability.  The  third  or  vol- 
umetric method  allows  of  a  balance  of  the  inequalities  of  size, 
and  best  portrays  "the  ratios  each  element  bears  to  the  others." 

By  the  numerical  method,  fifty  ants  would  be  placed  against, 
say,  six  ground-beetles.  A  computation  made  by  the  percentage- 
by-volume  method  would  doubtless  show  that  these  two  kinds  of 
food  represented  only  three  and  twenty-six  per  cent,  respectively, 
of  the  whole  food.  Hence  the  idea  furnished  in  the  first  case 
(a  ratio  of  50  to  6)  is  a  misleading  one.  Numbers  of  one  insect 
cannot  be  balanced  against  the  numbers  of  another  insect.  As 
each  bird  of  the  same  species  has  a  certain  average  stomach 
capacity,  the  ratio  of  each  element  to  this  average  capacity  gives 
the  most  accurate  idea  of  the  relative  proportions  of  each  kind 
of  food. 

Although  the  first  and  last  methods  have  been  those  most 
often  used  heretofore,  the  second  presents  certain  advantages 
(e.g.,  mathematical  accuracy)  which  should  not  be  overlooked. 
A  combination  of  the  numerical  and  the  percentage-by-volume 
method  has  been  used  in  the  present  work.  The  number  of  birds 
taking  the  different  kinds  of  food  offers  further  evidence  as  to 
their  capacity  for  good  or  ill. 

McAtee  (1912)  has  pointed  out  that  statements  of  numbers 
of  individuals  in  stomachs  has  an  interest  in  direct  proportion 
to  the  bigness  of  the  number.  Believing  this  to  be  largely  true, 
the  maximum  number  of  individuals  found  in  the  stomachs  has 
here  received  emphasis.  These  maximum  numbers  should  not 
be  considered  as  averages. 


1914]   Bryant:  Economic  Status  of  the  Western  Meadowlark     441 

It  does  not  seem  fair  to  compare  the  food  of  a  species  of 
bird  of  small  size  with  that  of  one  of  large  size  without  taking 
into  account  the  bird's  capacity.  The  degree  of  injury  or  benefit 
is  largely  dependent  on  the  total  amount  consumed.  In  this 
investigation  measurements  in  cubic  centimeters  of  the  stomach 
contents  of  a  number  of  western  meadowlarks  have  allowed  the 
computation  of  the  average  stomach  capacity.  For  male  birds 
this  average  capacity  is  three  cubic  centimeters ;  for  female  birds 
it  is  two  and  one-half  cubic  centimeters.  The  average  capacity 
is  two  and  three-fourths  cubic  centimeters. 

This  method  allows  of  some  interesting  computations  of  the 
capacity  for  good  or  harm  of  the  western  meadowlark.  If  there 
are  twelve  meadowlarks  to  the  square  mile,  as  there  are  in  many 
places,  these  birds  demand  over  one  hundred  cubic  centimeters 
of  food  daily.  This  capacity,  estimated  in  grain,  means  2200 
kernels,  in  grasshoppers  150  average-sized  individuals,  in  cut- 
worms 125  individuals,  in  ground-beetles  300,  in  ants  2500.  As 
digestion  is  constantly  going  on,  much  larger  numbers  are  in 
reality  taken,  as  is  evidenced  by  a  count  of  the  insects  in  a 
stomach.  These  numbers,  therefore,  are  not  even  a  minimum. 

As  pointed  out  (p.  412),  the  results  of  the  experiments  showed 
that  meadowlarks  must  completely  digest  a  meal  inside  of  four 
hours.  It  was  also  found  that  grain  takes  longer  to  digest  than 
do  insects.  Thus  it  can  be  seen  that  the  food  found  in  the  stomach 
at  any  one  time  does  not  represent  the  total  amount  of  food  taken 
daily,  but  only  about  a  third  part  of  that  consumed  daily.  Owing 
to  the  slower  digestion  of  grain,  the  amount  found  in  a  stomach 
must  represent  more  nearly  one-half  of  the  daily  requirement. 

Taking  an  average  capacity  of  two  and  three-quarters  cubic 
centimeters  and  considering  that  each  bird  nils  its  stomach  at 
least  three  times  a  day,  one  hundred  western  meadowlarks  must 
consume  near  a  liter,  or  about  a  quart  of  food  each  day.  If  the 
food  be  grain  it  can  be  seen  that  the  amount  of  destruction  is 
considerable ;  if  the  food  be  insects  it  can  be  seen  that  meadow- 
larks  take  a  large  daily  toll  of  insects.  The  same  type  of  com- 
putation shows  that  a  single  western  meadowlark  must  consume 
six  pounds  of  food  a  year. 


442  University  of  California  Publications  in  Zoology    [VOL.  11 

Some  idea  of  the  average  numbers  of  the  common  insects, 
grain,  and  weed  seeds  destroyed  at  each  meal  by  western  meadow- 
larks  can  be  obtained  from  the  following  tables,  which  were 
computed  from  the  results  of  stomach  examination  of  birds  col- 
lected in  Sacramento  and  Stockton,  California.  The  average  for 
the  day  is  three  times  that  for  each  meal. 


NUMBER  OF  COMMON  INSECTS,  GRAIN  AND  WEED  SEEDS  DESTROYED  BY 
WESTERN  MEADOWLARKS  COLLECTED  AT  SACRAMENTO,  CALIFORNIA 

February  to  April,  inclusive 


Grain 
1 

Weed 
seeds 

61 

( 

Beetles  ' 
6 

Cutworms 
and  cater-   Grass- 
pillars     hoppers 
8 

Ants, 
bees,  and 
Bugs         wasps     Spidei 

2 

71 

3             3 

6 

5 

4 
5 

1 

10 

4 

12 

1 

6 

8 

7 

5 

8 

3 

1 

9             2 

2 

5 

16 

10 

7 

11 

3 

5 

3 

12         .     . 

3 

13 

3 

2 

14 

7 

2 

Totals              5 

80 

56 

113 

1 

Av.  i)er  bird     3 

6 

4 

8 

9 

10 

11 

Totals 


Grain 


25 


28 


September  to  November,  inclusive 


Weed 
seeds 

60 
61 

155 

65 

37 

3 

200 
93 
80 

180 
15 

949 


Cutworm 

s 

Ants, 

and  catei 

•-   Grass- 

bees, anc 

I 

Beetles 

pillars 

hoppers 

Bugs 

wasps 

Spiders 

2 

1 

1 

1 

1 

1 

2 

1 

4 



2 

2 

6 

1 

8 

22 

2 

5 

1 

11 

2 

1 



2 

2 

2 
1 

22 


25 


10 
30 


1 

1 

12 


Av.  per  bird     2.5        86.2 


2.2 


2.6 


.6 


1914]   Bryant:  Economic  Status  of  the  Western  Meadowlark     443 

NUMBER  OF  INSECTS  TAKEN  BY  WESTERN  MEADOWLARKS  AT  STOCKTON,  SAN  JOAQUIN 

COUNTY,  CALIFORNIA 


ii  .s  i  i  1 1,  i  §  s .  i  i 

I2                  Date  J  £  I  g  J*  6  I  ff  3  1          I 

15     March-June,  1912  6  25  88  52  88  68  4  7  4  311        31 

Av.  per  bird                    .4       1.6       5.8       3.4  5.8       4.5  .2       .4       .2       20          2.0 

Av.  per  bird  per  day  1.2       4.8  17.4  10.2  17.4  13.5  .6  1.2       .6       60.0       6.0 


Few  people  have  any  realization  of  the  great  quantities  of 
insects  consumed  by  birds.  For  instance,  if  we  consider  that 
there  is  an  average  of  one  meadowlark  to  every  two  acres  of 
available  land  for  cultivation  (11,000,000  acres)  in  the  Sacra- 
mento and  San  Joaquin  valleys  and  that  each  pair  of  birds  raises 
an  average  of  four  young,  each  one  of  which  averages  one  ounce 
in  weight  while  in  the  nest  and  consumes  half  its  own  weight  of 
food  each  day,  it  takes  over  343 y%  tons  of  insect  food  each  day 
to  feed  the  young  birds  in  the  great  valleys  alone.  The  increased 
consumption  of  insect  food  due  to  nestling  birds  comes  at  a  time 
when  insects  are  most  numerous,  and  so  is  instrumental  in  helping 
to  prevent  an  undue  increase  of  insects.  As  insects  become 
injurious  only  when  in  maximum  numbers,  this  increased  con- 
sumption by  birds  is  doubly  important. 

A  conservative  estimate  of  the  approximate  amount  of  the 
different  kinds  of  food  consumed  by  the  average  meadowlark  in 
California  during  a  year  is  as  follows: 

Grain   1%  Ibs. 

Weed  seed    V2 

Insects  2% 

Total    6 

The  fact  that  the  western  meadowlark  eats  both  animal  and 
vegetable  food  is  a  point  in  its  favor.  If  it  were  exclusively 
insectivorous  the  bird  could  not  exist  in  such  large  numbers 
because  of  the  lack  of  insect  food  during  part  of  the  year.  The 
consequent  destruction  of  insect  life  would  therefore  be  much 
smaller. 


444          University  of  California  Publications  in  Zoology    [VOL.  11 

Certain  species  of  birds  when  hungry  will  not  only  fill  the 
stomach,  but  will  continue  eating  until  the  gullet  is  also  filled. 
This  is  often  found  to  be  the  case  with  linnets  and  bicolored 
blackbirds.  In  no  case,  however,  has  the  gullet  of  a  western 
meadowlark  been  found  well  filled  with  food.  At  the  most,  the 
last  insect  taken  before  the  bird  was  collected  has  been  found  in 
the  lower  part  of  the  gullet. 


CAPACITY  FOR  GOOD  OR  EVIL  AS  EVIDENCED  BY  THE  NUMBER  OF 
BIRDS  TAKING  THE  DIFFERENT  ITEMS  OF  FOOD 

The  percentage-volume  method  of  estimating  the  proportion 
of  the  different  kinds  of  food  taken  by  a  bird  gives  us  the  best 
idea  of  the  relative  importance  of  the  different  kinds  of  food 
in  the  diet  of  a  given  bird.  However,  the  frequency  of  occur- 
rence of  the  different  items  in  the  food,  shown  by  a  statement 
of  the  number  of  birds  taking  each  item,  furnishes  additional 
evidence  as  to  the  capacity  of  a  species  for  good  or  ill.  The 
number  of  birds  taking  a  certain  kind  of  insect  food  can  be 
regarded  as  an  approximate  index  of  the  availability  of  that 
kind  of  food,  and  to  a  much  less  extent  as  an  index  of  food 
preference  if  we  consider  insects  as  being  evenly  distributed  and 
birds  as  being  but  slightly  influenced  by  psychological  processes. 
A  nearer  approximation  can  be  obtained  by  multiplying  the 
number  of  birds  by  the  number  of  insects  taken.  In  such  a 
computation  the  number  of  insects  taken  is  considered,  as  well 
as  the  number  of  birds  taking  the  different  elements  of  food. 
By  this  method  the  index  of  availability  of  crickets  at  Live  Oak, 
Sutter  County,  was  four  in  1911,  whereas  this  index  was  6902 
at  Hollister,  San  Benito  County,  in  the  same  year.  By  the  same 
method  of  calculation  the  index  of  availability  of  grasshoppers 
was  2162  at  Live  Oak  and  but  1541  at  Hollister.  In  the  first 
case  crickets  were  1670  times  as  available  at  Hollister  as  at 
Live  Oak,  and  grasshoppers  1.4  times  more  available  at  Live  Oak 
than  at  Hollister.  The  following  table  gives  a  comparison  of 
availability  of  the  commoner  insects  as  evidenced  by  indices  of 
availability.  Preference  is  here  classed  as  a  factor  in  availability. 


1914]   Bryant:  Economic  Status  of  the  Western  Meadowlark     445 


01   01   en  Number  of 


o  o   o   o     birds 

^  ^  ^>  ^>                                 j 

4         ^ 

2    & 
>           a    g 

^  t^ 

CD      ^ 

CD      CD     O     CS                                              / 

H     >-i     <rt-    <n-                                      J; 

3          §   ^ 

H      O' 

CD     CD      M      M                                              J 

s-  s 

1     * 

UH 

g-  5-  a  B                    ! 

=       "8  S, 

QTQ     O 

i—  i 

00    (JQ     §     §                                         * 

^                0     53* 

o    cr1 

1 

8S||                     1 

i     ?§; 

3           S»  5 

H,     -' 

2t  % 

3 

sr1  IT  3  S                        ' 

"-« 
1           PJ  ^ 

&  ^ 

H 

§  S  IT  S3"                       ' 

a           "  pr 

<rt-     t-i- 

t^ 

g 

<^§S                       ! 

S           ^  tw 

P      3 

^-  P    a4  a- 

J,        (»' 

(jq     I 

1 

3 

H 

§       ^    'tf*     0, 

3        j 

i     i 

5T         m    p. 

2j 

> 

•3                     oo 

O5 

to 

—  •    x 

i     *     8    . 

i            ^    : 

^             rfx        Wireworms 

^  o  Oats 

w 

02 

ft                        IN3 

2             co  co  Bugs 

^ 

^j 

•^           (Hemiptera) 

j 

^   01  Barley 

g 

g  g         M  Grain 

M 

i 

O     GO 

•      ^  Cicadas 

i—  i 

o 

?3 

—  ^            —i 

i_i 

.     .     ^  01  Weed  seeds 

•J 

^.                           CO 

01  to  Wheat 

bo 

Si 

> 

CO    CO 

^                  h-  '    tO 

o 

-^ 

^             05   to  Ants 

O 

to  to 
o  ^  5  o  Beetles 

l*q     QQ 

•^ 

1 
So  ^ees  an(^ 

co        Corn 

oo 
J^        Sorghum 

3 
cr 
CD 

CD 

1 

tO    M 

to          wasps 

H 

^z 

g  g  °*  ^Crickets 

25 

^                   to 

01  „ 

r,.        Beans 

S 

1 

72                    __*     ^j|  ^.    . 

H-  '• 

H-i 

to  to 

O                   CO 

P 

(Jl 

CO     h^     O5     ^ 

oo   01  co  o  Grasshoppers 

^  gg  Weed  seeds 

P^ 

o 

bo  bi 

5            V         Butterflies 

• 

i—i 

^ 

8 

r^ 

W 

>_.  j_i  co  M  Jerusalem 

§                          -q 

,_i 

o 

» 

tc   oo                crickets 

i_5            co  05  pupae 

•     °°  Grape  seeds 

72 

h-1             CO    Ol 

^i  «o  to  o  Cutworms  and 

Tl                                               CO 

to 

H3 

O5  to                caterpillars 

.^  w  Flies 

r  w  Grass 

CO 

5 

^1    00 

01  oo  H-  oBugs 

Cd 

1—  » 

o 

O    CO 

*"            La   ^  Fly  larvae 

O 

01  MX  Beetles 

d 

LCl  i_i  E3  M  Bees  and 

es 

•H                                          (^7| 

to 

2 

o  rf^               wasps 

^             to        Crane  flies 

to 

g 

^    -3     *. 

^                   ~^ 

N*   W  _  .  . 

to  co  Crickets 

w 

C5  bo             Ants 

?°   ^  Spiders 

J^. 

^ 

t»  i            "^i 

W 

e3 

Ol 

r^ 

50  Spiders 

^             M  01  Spider  egg 
co          cases 

o  2q  Grasshoppers 

01 

C 
O2 

a  co 
J->  bs             Miscellaneous 

^                          05 

g            co        Centipeds 

u,  §  Jerusalem 
K.J          crickets 

O 

•=a 

^                   oo 

o 

rf^        Millipeds 

Ol 

o 

01  O5              Animal 
^,  M               food 

^             .      °  Sow  bugs 

^  ^  Cutworms  and 
•             caterpillars 

W 

W 

4-  co              Vegetable 

00 

co  co                food 

446          University  of  California  Publications  in  Zoology    [VOL. 11 
INDICES  OF  AVAILABILITY  OF  COMMON  INSECTS  EATEN  BY  WESTERN  MEADOWLARKS 

No.  of  Cut-  Grass-  Bees  and 

Locality                              birds  Beetles  worms  hoppers  Crickets  Bugs  wasps  Ants 

Live  Oak,  gutter  Co.           60  1,775  288  2,162  4  533  78  18 

Hollister,  San  Benita  Co.     60  4,307             7  1,492  6,680  197  265         35 


The  most  available  and  the  most  popular  food  of  the  western 
meadowlark,  if  it  may  be  judged  by  the  frequency  with  which 
it  has  been  found  in  the  stomachs,  is  beetles.  Seventy-five  per 
cent  of  all  the  stomachs  examined  contained  beetles  (Coleoptera). 
Vegetable  food  in  the  form  of  oats  is  next  in  order  of  frequency, 
thirty-four  per  cent  of  the  meadowlarks  examined  having  taken 
oats  as  food.  Grasshoppers,  cutworms  and  caterpillars,  and  weed 
seeds  were  found  in  thirty  per  cent,  twenty-six  per  cent,  and 
twenty-five  per  cent  respectively  of  the  stomachs  examined. 

The  accompanying  table  gives  a  summary  of  the  number  of 
times  each  article  of  diet  was  taken  and  the  percentage  of  each 
article  in  the  diet  of  the  species.  Attention  should  be  called 
to  the  fact  that  a  very  large  percentage  (seventy  per  cent)  of 
the  birds  taking  oats  took  wild  oats  (Avena  fatua)  instead  of 
the  tame  varieties.  Consequently  not  more  than  ten  per  cent 
of  the  birds  examined  had  taken  cultivated  varieties  of  oats. 

Although  the  percentages  showing  the  proportionate  number 
of  times  each  kind  of  food  is  taken  to  the  number  of  birds  exam- 
ined differs  from  the  percentages  showing  the  proportionate  vol- 
ume of  each  kind  of  food,  yet  they  parallel  each  other  to  a 
considerable  extent.  The  accompanying  table,  giving  only  th^ 
principal  articles  of  diet,  shows  this  parallelism  in  the  per- 
centages. 


IV. — PERCENTAGE-VOLUME  AND  PERCENTAGE  OF  WESTERN  MEADOWLARKS  TAKING  DIFFERENT 

ELEMENTS  OF  FOOD 


1  II  *  ! 

03  03                                                          ,J3  CO    Q, 

11  I  S«  I      ii  35 

6             &  ~              6              5  £""  <d          pr  ^  t"" 

Percentage-volume  ...     30.1         9.9  17.0         3.5       18.0  3.1  3.0         4.0  .1 
Percentage  of  birds 

taking  46.0       25.0  75.2       12.4       30.5  13.4  16.7       19.2  1.1 


1914]   Bryant:  Economic  Status  of  the  Western  Meadowlark     447 

FOOD  OF  NESTLINGS 

Three  methods  are  available  for  determining  the  food  of 
nestlings ;  first,  the  rate  of  feeding  may  be  determined  by  watch- 
ing the  number  of  trips  made  to  the  nest  by  the  adult  birds  while 
feeding  the  young;  second,  the  young  birds  may  be  made  to 
disgorge  their  food;  or  third,  the  bird  may  be  killed  and  the 
food  in  the  digestive  tract  examined.  The  first  method  has  not 
been  used  with  any  great  degree  of  success  because  of  the  diffi- 
culty in  approaching  near  enough  to  the  nest  to  observe  the 
feeding.  The  western  meadowlark  does  not  become  accustomed 
to  an  intruder  so  easily  as  do  other  birds.  The  food  observed 
in  the  bills  of  adult  birds  carrying  food  to  their  young  has  most 
often  been  cutworms  or  grasshoppers.  One  female  bird,  while 
feeding,  persistently  flew  into  an  oat  field  and  returned  each 
time  with  cutworms.  Another  female  bird  was  seen  to  catch 
three  or  four  grasshoppers  in  her  bill,  and  then  fly  to  the  nest. 

Examination  of  the  digestive  tract  of  nestling  birds  has 
shown  that  they  are  fed  very  largely  on  cutworms,  grasshoppers, 
and  ground-beetles.  The  stomachs  of  two  nestlings  obtained  from 
the  same  nest  contained  egg-shells.  This  is  not  an  unusual  occur- 
rence, for  nestling  birds  of  other  species  are  fed  on  the  egg-shells. 
In  no  case  was  grain  found  in  the  stomachs,  and  in  only  a  few 
cases  were  weed  seeds  found.  Young  birds  doubtless  need  a 
larger  quantity  of  food  than  adults.  In  almost  every  case  the 
stomachs  were  average  full  or  over.  This  means  that  each  stomach 
contained  nearly  three  cubic  centimeters  of  food.  Since  it  has 
been  shown  that  young  birds  need  over  one-half  their  own  weight 
of  food  each  day,  the  birds  when  hatched  must  consume  about 
one-fourth  of  an  ounce  a  day,  and  when  ready  to  fly  about  two 
ounces. 

This  increased  consumption  of  insects  due  to  the  demands 
of  young  birds  comes  at  a  time  when  there  are  growing  crops 
which  need  protection  and  when  insects  are  most  numerous,  thus 
emphasizing  the  value  of  birds  as  balancers.  The  fact  that 
meadowlarks  show  a  greater  preference  for  certain  kinds  of  food 
while  feeding  the  young  enlarges  their  sphere  of  usefulness. 

In  spite  of  the  fact  that  a  certain  amount  of  grain  and  weed 
seed  is  available,  young  nestling  birds  are  fed  almost  entirely 


448          Un  iversity  of  California  Publications  in  Zoology    [VOL.  11 

on  insects.  It  would  seem,  therefore,  that  there  is  a  certain 
preference  for  insect  food  shown  at  this  time  of  the  year.  It  is 
a  well-known  fact  that  many  adult  birds  which  feed  on  weed 
seeds  feed  their  young  almost  entirely  on  insects.  It  seems  safe 
to  say,  therefore,  that  the  western  meadowlark,  when  feeding 
the  young,  turns  its  attention  to  insects. 

The  accompanying  table  (Table  V,  p.  445)  gives  a  comparison 
of  the  food  of  fifty  juvenile  and  fifty  adult  western  meadowlarks. 

VARIATION  IN  KIND  OF  FOOD 

The  kind  of  food  taken  by  the  western  meadowlark  is  de- 
pendent upon  two  cycles,  the  individual  cycle  and  the  environ- 
mental cycle. 

The  individual  cycle  includes  such  factors  as  individual  taste, 
time  of  feeding,  and  the  preference  for  a  particular  locality. 
In  fact,  all  of  the  factors  which  depend  upon  the  individual  tastes 
or  habits  of  the  bird  are  grouped  here.  Whether  an  individual 
bird  has  a  particular  taste  for  a  certain  insect,  it  is  impossible 
to  determine  without  experiment.  Since  we  find  different  indi- 
viduals showing  different  characteristics,  we  can  safely  infer  that 
each  individual  may  show  a  slight  preference  for  one  kind  of 
food  above  another.  Then,  too,  the  time  during  which  the  bird 
feeds  has  some  effect  upon  the  availability  of  certain  kinds  of 
food,  so  that  we  should  naturally  expect  that  the  kind  of  food 
would  be  governed  to  some  extent  by  the  time  the  bird  chooses 
for  feeding.  The  particular  locality  frequented  by  the  bird  must 
also  influence  the  kind  of  food  taken.  The  food  of  a  bird  feeding 
entirely  in  a  grain  field  would  certainly  show  a  slight  variation 
from  that  taken  by  one  living  entirely  in  a  pasture. 

The  environmental  cycle  takes  into  account  the  changes  in 
the  availability  of  insects  and  seeds,  due  to  seasonal  and  climatic 
conditions.  The  maximum  supply  of  weed  seed  is  available  dur- 
ing September  and  October.  The  maximum  supply  of  insects 
is  apparently  available  during  May  and  June.  Even  the  culti- 
vation of  land  has  much  to  do  with  the  availability  of  certain 
kinds  of  food.  Many  weed  seeds  are  easily  obtained  during 
September  and  October,  which,  after  plowing  begins,  are  hidden 


1914]   Bryant:  Economic  Status  of  the  Western  Meadoivlark     449 

beneath  the  soil  and  so  made  unavailable.  This  helps  to  explain 
why  birds  feed  so  largely  on  grain  during  the  winter  months. 
Grain  is  far  more  available  in  cultivated  districts  at  this  time 
than  weed  seed. 

In  some  localities  even  a  daily  cycle  may  be  noted.  Certain 
insects  appear  in  larger  numbers  after  the  sun  has  warmed  up 
the  soil.  Certain  insects,  such  as  cutworms,  are  found  in  greater 
abundance  before  sunrise.  Thus  we  might  naturally  expect  that 
birds  collected  early  in  the  morning  would  have  taken  a  larger 
percentage  of  cutworms  than  birds  collected  later  in  the  day. 
Stomach  examination  has  substantiated  this  as  a  fact. 

The  kinds  of  crops  raised  in  any  particular  locality  must 
also  influence  the  kinds  of  food  taken,  for  it  changes  their  avail- 
ability. For  instance,  the  stomachs  of  several  meadowlarks  con- 
tained Egyptian  corn  and  milo  maize.  Where  this  crop  is  raised 
to  any  extent  the  meadowlark  doubtless  occasionally  turns  its 
attention  to  this  type  of  grain.  In  other  localities  where  only 
oats  and  barley  are  grown  these  grains  are  the  only  ones  available. 


VI. — PERCENTAGES  OF  FOOD  OF  MEADOWLARKS  TAKEN  IN  ALFALFA  FIELDS  IN  THE 
VICINITY  OF  HANFORD,  CALIFORNIA 


Miscellaneous 

1  potato-bug 

1  potato-bug,  1  bee 

1  potato-bug 
1  fly,  3  ants 

10  ants 
1  fly 

1  pupa 


17  small  flies 

1  ant 

1  cricket,  2  ant-lions 

16.1 


Number 

Per  cent 

Animal 
matter 

Vegetable 
matter 

cut- 
worms 

cut- 
worms 

Number 
beetles 

Per  cent 
beetles 

1 

100 

3 

50 

4 

15 

2 

96 

4 

2 

30 

7 

46 

3 

100 



25 

80 

9 

20 

4 

98 

2 

1 

6 

5 

100 

5 

65 

2 

12 

6 

96 

4 

19 

76 

6 

20 

7 

100 



9 

60 

7 

20 

8 

88 

12 

23 

75 

4 

10 

9 

85 

15 

1 

10 

7 

75 

10 

94 

6 

21 

83 

2 

8 

11 

100 

2 

80 

2 

20 

12 

100 



I 

16 

10 

76 

13 

100 

4 

55 

6 

20 

14 

100 



4 

85 

3 

15 

15 

17 

83 

2 

10 

1 

4 

16 

100 

8 

60 

5 

14 

Totals  and 

averages 

92.1 

7.9 

129 

52.1 

76 

24.3 

450          University  of  California  Publications  in  Zoology    ["^OL.  11 

Stomach  examination  has  clearly  demonstrated  the  fact  that 
birds  collected  in  alfalfa  fields  consume  much  larger  quantities 
of  cutworms  and  caterpillars  than  birds  collected  in  grain  fields 
in  the  same  general  locality.  The  following  table  computed  after 
the  examination  of  sixteen  stomachs  of  meadowlarks  collected 
in  alfalfa  fields  in  the  vicinity  of  Hanford,  Kings  County.  Cali- 
fornia, clearly  brings  out  the  increased  percentage  of  cutworms 
and  caterpillars  in  the  food  taken  by,  birds  in  such  fields.  An- 
other table  showing  a  comparison  of  the  food  of  thirty-four  birds 
collected  in  the  above  locality,  but  in  different  kinds  of  fields, 
brings  out  the  same  point. 

VII. — COMPARISON  OF  FOOD  TAKEN  BY  WESTERN  MEADOWLARKS  COLLECTED 
IN  ALFALFA  FIELDS,  GRAIN  FIELDS,  ORCHARDS,  AND  VINEYARDS 

Averages  of  nine  birds  per  month  collected  in  March,  April,  and  May,  1911,  at 
Hanford,  Kings  County,  California 


Average 

number  of 

Per  cent 

Per  cent 

Per  cent 

Average 

cutworms 

cutworms 

Kind  of 

animal 

vegetable 

number  of 

Per  cent 

and  cater- 

and cater- 

field 

food 

food 

beetles 

beetles 

pillars 

pillars 

Alfalfa 

95.9 

4.1 

5.1 

24.1 

11.2 

61.4 

Grain 

99.4 

.6 

9.4 

58.5 

1.8 

21.0 

Orchard 

98.3 

1.7 

4.8 

31.7 

6.5 

55.7 

Vineyard 

78.3 

21.7 

.1 

5.0 

7.0 

42.0 

VARIATION  OF  FOOD  ACCORDING  TO  TIME  OF  YEAR 

Abundant  data  have  allowed  a  comparison  of  the  food  of  the 
western  meadowlark  by  the  hour,  day,  week,  month,  and  year. 
Little  change  in  kind  of  food  can  be  noted  from  one  time  of  day 
to  another  unless  birds  were  taken  in  different  localities.  Never- 
theless, the  quantity  of  food  found  in  the  stomachs  varies  with 
the  time  of  day.  The  maximum  is  found  about  nine  o'clock  in 
the  morning,  and  the  minimum  from  one  to  two  o'clock  in  the 
afternoon.  The  same  can  be  said  of  the  food  from  day  to  day. 
A  comparison  of  the  food  from  week  to  week,  however,  shows 
considerable  change. 

The  change  from  week  to  week  (and  the  same  can  be  said 
of  the  change  from  month  to  month)  closely  parallels  the  avail- 
ability of  the  different  articles  of  diet.  Weed  seed  and  waste 
grain  are  nearly  always  available  during  September  or  October. 


1914]   Bryant:  Economic  Status  of  the  Western  Meadowlark     451 

Yet  if  enough  grasshoppers  and  ground-beetles  are  available,  the 
birds  evidently  take  these  in  preference,  for  stomachs  are  more 
often  found  filled  with  insects  than  with  weed  seeds.  The 
accompanying  diagrams  illustrate  the  great  change  in  food  habits 
from  one  time  of  year  to  another.  It  will  be  noted  in  each 
instance  that  the  food  of  the  western  meadowlark  is  made  up 
largely  of  insects  during  the  spring  and  summer  months  and 
largely  of  grain  and  weed  seeds  during  the  fall  and  winter 
months.  It  is  needless  to  point  out  that  this  parallels  the  avail- 
ability of  insect  food,  and  to  a  less  extent  the  availability  of 
vegetable  food.  In  that  the  percentage  of  animal  food  for  the 
year  is  greater  than  the  percentage  of  vegetable  food,  and  since 
some  insect  is  nearly  always  found  in  stomachs  filled  with  grain 


Animal  food 


Vegetable  food 


Jan. 
Feb. 
Mar. 
Apr. 
May 
June 
July 
Aug. 
Sept. 
Oct. 
Nov. 
Dec. 


Total 


Fig.  C. — Diagram  showing  change  of  food  habits  of  the  western  mead- 
owlark from  month  to  month.  Note  that  the  maximum  consumption  of 
animal  food  is  to  be  found  in  May,  June,  and  July,  and  the  minimum, 
corresponding  with  the  maximum  of  vegetable  food,  in  January  and 
February.  Computed  from  the  results  of  stomach  examinations  of  an 
average  of  twelve  birds  taken  each  month  of  the  year  at  Ked  Bluff, 
Tehama  County,  California. 


and  weed  seeds,  whereas  grain  and  weed  seeds  are  far  less  often 
found  in  stomachs  filled  with  insects,  it  seems  safe  to  say  that 
animal  food  is  preferred  and  vegetable  food  is  used  as  a  make- 
shift. 

The  diagram  showing  the  food  of  western  meadowlarks  col- 
lected in  the  vicinity  of  Red  Bluff  (fig.  C)  illustrates  this  pref- 
erence for  insects.  Ninety  per  cent  of  the  birds  examined  were 


452  University  of  California  Publications  in  Zoology    [VOL.  11 

collected  in  grain  fields,  and  yet  the  only  time  when  they  turned 
to  grain  was  in  the  winter,  when  the  numbers  of  insects  were  at  a 
minimum.  Evidently  grain  fields  furnish  an  abundant  supply 
of  animal  food  during  most  of  the  year.  The  maximum  con- 
sumption of  animal  food  is  in  June  and  the  minimum  in  January. 
The  minimum  of  animal  food  corresponds  necessarily  to  the 
maximum  in  vegetable  food. 

The  diagram  showing  the  proportion  of  the  two  kinds  of  food 
of  birds  taken  in  the  vicinity  of  San  Bernardino  (fig.  D)  lacks 
some  of  the  inaccuracies  to  be  noted  in  the  former  diagram.  The 
maximum  consumption  of  animal  food  is  in  April  instead  of 
May,  due  to  the  difference  in  climatic  conditions.  The  birds 
taken  in  this  locality,  although  collected  largely  in  grain  fields, 
consumed  an  unusually  large  proportion  of  animal  food  during 
1911,  more,  in  fact,  than  any  other  series  of  birds  collected  in 
southern  California. 

The  amounts  of  food  taken  by  meadowlarks  collected  every 
two  weeks  at  Newman,  Stanislaus  County  (fig.  E),  also  clearly 
brings  out  the  change  in  food  habits  from  one  part  of  the  year  to 
another. 

The  averages  of  the  different  kinds  of  food  for  the  year  do 
not  change  greatly  from  one  year  to  another.  The  time  of  year 
when  the  different  kinds  of  food  reach  a  maximum  in  the  diet 


Fig.  D. — Diagram  showing  change  of  food  habits  of  the  western  mead- 
owlark  from  month  to  month.  Note  that  the  maximum  consumption  of 
animal  food  is  in  April  and  May.  Computed  from  stomach  examinations 
of  an  average  of  six  birds  collected  each  month  in  a  year  at  San  Bernar- 
dino, San  Bernardino  County,  California. 


1914]   Bryant:  Economic  Status  of  the  Western  Meadowlark     453 

does  change.  This  seems  natural,  for  weather  conditions  affect 
the  available  food  supply.  Thus  we  find  very  few  ants  taken 
by  meadowlarks  collected  at  Newman,  Stanislaus  County,  in  1911, 
but  large  numbers  taken  during  the  same  months  in  1912  (fig.  E). 


Animal  food 


Vegetable  food 


Fig.  E. — Diagram  showing  the  comparative  amounts  of  the  different 
kinds  of  food  of  the  western  meadowlark  every  two  weeks  during  a  year. 
Computed  from  the  results  of  stomach  examinations  of  birds  collected  at 
Newman,  Stanislaus  County,  California. 


The  following  table  gives  a  comparison  of  the  amounts  of 
animal  and  vegetable  food  and  the  amounts  of  some  of  the 
common  elements  of  food  taken  by  meadowlarks  at  Newman, 
Stanislaus  County,  in  1911  and  in  1912.  The  difference  in 
amounts  of  animal  and  vegetable  food  is  seven  per  cent.  A  large 
increase  in  1912  of  the  number  of  grasshoppers  taken  can  be 
noted.  Seventy-two  birds  from  each  year  were  examined. 


COMPARISON  OF  FOOD  OF  MEADOWLARK  IFOR  Two  SUCCESSIVE  YEARS 


Year 
1911 
1912 


Per  cent 

Per  cent 

Per  cent 

animal 

vegetable 

Per  cent 

grass- 

Per cent 

Per  cent 

food 

food 

grain 

hoppers 

beetles 

ants 

53.5 

46.5 

38.4 

27.5 

8.5 

.4 

65.7 

34.3 

32.3 

42.4 

9.9 

3.0 

454          University  of  California  Publications  in  Zoology    [VOL.  11 

VARIATION  OF  FOOD  HABITS  ACCORDING  TO  LOCALITY 
The  comparison  of  the  food  of  the  western  meadowlark  in 
California  has  been  made  in  several  different  ways.  As  soon  as 
it  was  found  definitely  that  the  food  varied  greatly  from  one 
part  of  the  state  to  the  other,  it  was  decided  to  make  three  group- 
ings for  comparison  as  follows:  (1)  districts;  (2)  counties;  (3) 
localities.  The  wide  difference  in  the  kind  and  availability  of 
food  in  the  coast  region  and  in  the  interior  valleys  makes  this 
comparison  of  prime  importance.  Although  a  comparison  by 
counties  covers  in  a  measure  that  by  districts,  yet  this  smaller 
unit  of  area  brings  to  each  rancher  a  knowledge  of  the  food  of 
the  meadowlark  in  his  own  county.  Although  the  difference  in 
locality  will  parallel  largely  the  difference  in  county,  owing  to 
the  fact  that  usually  but  one  series  of  birds  was  collected  in  each 
county,  yet  this  comparison  affords  a  knowledge  as  to  the  amount 
of  variation  between  still  smaller  units  of  area. 

Birds  collected  in  the  vicinity  of  San  Diego  took  a  smaller 
percentage  of  insects  than  those  collected  at  Riverside  and  San 
Bernardino.  Birds  collected  in  the  vicinity  of  Ukiah,  Mendocino 
County,  took  larger  numbers  of  click-beetles  than  birds  from 
any  other  locality.  Stomachs  of  birds  collected  in  the  vicinity 
of  Hollister,  San  Benito  County,  contained  extraordinarily  large 
quantities  of  crickets.  At  Hanford,  Kings  County,  birds  fed 
very  extensively  on  cutworms.  Birds  collected  at  Newman,  Stan- 
islaus County,  contained  extra  large  percentages  of  stink-bugs 
and  grasshoppers.  Birds  from  Red  Bluff,  Tehama  County,  Live 
Oak,  Yuba  County,  Sacramento,  Sacramento  County,  Newman, 
Stanislaus  County,  and  Los  Banos,  Merced  County,  took  very 
nearly  the  same  proportion  of  animal  and  vegetable  food. 

VARIATION  OF  FOOD  ACCORDING  TO  DISTRICTS 
For  the  purpose  of  comparison  the  following  county  groupings 
were  made: 

Humid  Coast  Belt:      Humboldt,  Mendocino,  Sonoma,  Marin. 

Interior  Valleys:          Tehama,  Butter,  Sacramento,  San  Joaquin,  Alameda, 

Merced,  Stanislaus,  Madera,  Fresno,  Kern,  Kings. 
Mountain  District:      Shasta,  Lassen,  Nevada,  Calaveras. 
Arid  Coast  Belt:  Ventura,   Los   Angeles,    San   Bernardino,    Kiverside, 

Orange,  San  Diego. 
Desert:  Inyo,  Imperial. 


1914]   Bryant:  Economic  Status  of  the  Western  Meadowlark     455 

The  following  table  gives  the  food  of  the  western  meadowlark 
in  these  districts : 

TABLE  SHOWING  VARIATION  OF  POOD  ACCORDING  TO  DISTRICTS 


Humid  Coast  Belt 

Number 
of 
birds 

99 

Per  cent 
animal 
food 

59.0 

Per  cent 
vegetable 
food 

41.0 

Interior  valleys  

785 

61.8 

38.2 

Mountain  district 

27 

69  6 

30  4 

Arid  Coast  Belt 

536 

58.5 

41  5 

Desert  ... 

58 

63.0 

37.0 

By  considering  only  those  localities  where  complete  or  nearly 
complete  series  were  obtained,  a  still  more  marked  difference 
could  be  noted.  If  the  results  from  an  examination  of  more 
birds  from  the  humid  coast  belt  and  mountain  districts  were 
available,  the  results  would  show  a  larger  amount  of  animal  food. 
The  remarkable  thing  is  that,  considering  the  widely  different 
climatic  conditions,  the  food  averages  do  not  vary  more  than  ten 
per  cent. 

INFLUENCE  OF  AGE  AND  SEX  ON  QUANTITY  OF  FOOD  TAKEN 

As  the  male  weighs  an  ounce  (28.35  grams)  more  than  does 
the  female,  a  difference  of  one-half  cubic  centimeter  in  capacity 
is  accounted  for.  Juveniles  average  less  food  than  adults.  This 
is  explainable  on  the  ground  that  they  are  less  experienced  in 
obtaining  food.  Male  and  female  nestlings  apparently  have  more 
nearly  the  same  capacity  than  do  adults  of  different  sexes. 

A  slight  variation  has  been  noted  between  the  food  of  adults 
and  young  birds,  both  in  kind  and  quantity.  Young  birds  ap- 
parently lack  the  experience  of  the  adults  and  pick  up  only  the 
more  conspicuous  insects  and  weed  seeds.  They  also  lack  the 
experience  needed  for  catching  certain  insects,  and  therefore  the 
stomachs  average  a  little  less  in  volume  of  food.  It  has  been 
pointed  out  by  Finn  (1887)  "that  each  bird  has  to  separately 
acquire  its  experience,  and  it  well  remembers  what  it  has 
learned."  There  appears  to  be  little  instinctive  knowledge  of 
the  different  kinds  of  food,  and  each  young  bird  must  test  and 
learn.  It  would  seem,  therefore,  that  young  birds  would  collect 
many  so-called  protected  insects,  whereas  some  experienced  ones 


456          University  of  California  Publications  in  Zoology    [VOL.  11 

would  pass  them  by.  This  theory  has  not  been  supported  by 
the  results  of  this  investigation,  nor,  on  the  other  hand,  has  it 
been  broken  down.  Although  young  birds  were  distinguished 
from  adults,  yet  the  unusual  insects  found  were  not  constantly 
taken  from  the  stomachs  of  young  birds. 


COMBINATION  OF  FIELD  AND  LABORATORY  WORK 

Judd  (1901)  has  suggested,  as  the  best  means  of  determining 
the  food  of  birds,  an  examination  of  the  available  food  supply 
combined  with  stomach  examinations  of  birds  taken  in  the  same 
Iocalit3r.  There  are  many  points  which  commend  this  method. 
Yet  there  is  great  difficulty  in  determining  the  available  food 
supply.  Even  an  experienced  observer  cannot  estimate  even 
with  moderate  accuracy  the  comparative  numbers  of  insects  and 
weed  seeds  in  any  given  locality.  A  bird  is  able  to  see  many 
articles  of  diet  which  such  an  observer  doubtless  overlooks.  Let 
me  cite  a  case  in  point :  A  meadowlark  was  seen  to  fly  to  a  grain 
field  and  collect  cutworms  to  feed  its  young.  Careful  investi- 
gation by  me  of  the  place  where  the  cutworms  were  collected 
failed  to  reveal  any.  The  same  was  noted  with  a  pair  of  Brewer 
blackbirds  who  persistently  collected  cutworms  in  a  pasture. 
Continued  investigation  in  the  same  pasture  did  not  allow  of 
the  collection  of  a  single  cutworm. 

An  attempt  was  made  to  follow  this  particular  line  of  investi- 
gation, but  was  finally  given  up  on  account  of  the  multitude  of 
personal  errors  which  are  easily  introduced.  It  seemed  best  to 
concentrate  on  the  usual  method  of  stomach  examination,  thereby 
making  comparison  with  previous  work  possible. 


RELATION  OF  BIRDS  TO  INSECT  OUTBREAKS 

The  value  of  birds  as  insect  destroyers  is  more  noticeable  at 
the  time  of  an  insect  outbreak.  Their  importance  in  maintaining 
an  equilibrium  depends  largely  upon  their  effect  when  insects 
occur  in  abnormal  numbers  and  thus  become  noticeably  injurious. 
In  two  insect  outbreaks  investigated  the  western  meadowlark  was 
found  to  take  a  very  active  part  in  insect  destruction. 


1914]   Bryant:  Economic  Status  of  the  Western  Meadowlark     457 

During  the  spring  and  summer  of  1911  the  nymphalid  butter- 
fly Eugonia  calif  ornica  became  very  abundant  in  the  northern 
part  of  the  state.  Since  butterflies  are  seldom  eaten  by  birds, 
the  outbreak  afforded  a  splendid  opportunity  to  study  the  food 
habits  of  birds.  Consequently  an  investigation  was  carried  on 
during  the  latter  part  of  August  at  Sisson,  Siskiyou  County, 
California,  when  these  insects  were  in  abnormal  numbers. 

Apparently  because  of  the  availability  of  the  insect,  several 
birds  were  found  to  be  destroying  butterflies.  The  Brewer  black- 
bird (Euphagus  cyanocephalus)  fed  on  them  almost  exclusively, 
whereas  the  western  meadowlark  (StumeUa  neglect  a),  western 
kingbird  (Tyrannus  verticalis),  blue-fronted  jay  (Cyanocitta 
stellari  frontalis),  and  Say  phoebe  (Sayornis  say  us)  took  them 
sparingly.  A  comparison  of  the  food  of  birds  taken  before  the 
plague  with  that  of  birds  taken  while  the  plague  was  at  its  height 
showed  that  birds  had  varied  their  food  habits  and  had  taken 
advantage  of  the  abundant  supply  of  insect  food  in  the  form  of 
butterflies. 

Both  observation  and  stomach  examination  showed  the  western 
meadowlark  to  feed  on  the  butterfly  (Eugonia  californica).  A 
lone  meadowlark  feeding  with  some  Brewer  blackbirds  on  the 
grass  plot  adjoining  the  station  at  Sisson  was  seen  to  run  after 
several  butterflies  and  to  catch  one.  In  the  examination  of  seven 
stomachs,  two  contained  butterflies.  Fifteen  and  two-tenths  per 
cent  of  the  food  taken  by  the  five  meadowlarks  collected  in 
August  was  made  up  of  butterflies.  All  of  these  birds  were 
taken  in  meadow  or  cut  fields  of  wild  hay  where  other  insect  life 
was  abundant.  Beetles  and  grasshoppers  formed  the  bulk  of 
the  food. 

Stronger  evidence  that  birds  turn  their  attention  to  the  insect 
most  available  can  hardly  be  found,  for  in  this  case  we  find  a 
supposedly  unpalatable  insect  becoming  food  for  a  number  of 
species  of  birds.  The  United  States  Biological  Survey,  in  the 
examination  of  more  than  40,000  stomachs,  has  found  but  four 
records  of  birds  eating  butterflies,  ''and  one  of  these  probably 
relates  to  the  capture  of  a  very  recently  emerged  specimen,  or 
to  one  torn  from  the  pupa  before  emergence,  as  it  was  accom- 
panied by  a  pupa  of  the  same  species."  (See  McAtee,  1912c.) 


458          University  of  California  Publications  in  Zoology    [VOL.  11 


Whether  butterflies  are  too  active  to  be  caught,  or  whether  they 
are  unpalatable  because  of  odor  or  taste,  are  questions  still  await- 
ing an  answer.  In  any  case  it  can  truly  be  said  that  butterflies, 
considering  their  abundance,  are  not  taken  as  food  in  anywhere 
near  the  proportion  that  other  insects  are  taken.  That  four  of 
the  larger  common  birds  of  the  region  should  have  fed  upon  these 
insects  and  that  one  of  these  should  have  fed  almost  entirely  upon 
them  is  certainly  significant. 

It  cannot  be  said  that  lack  of  other  food  caused  these  birds 
to  turn  their  attention  to  butterflies,  for  many  of  the  birds  col- 
lected had  either  taken  no  butterflies  or  but  one  or  two.  The 
butterflies  were  not  only  conspicuous,  but  extremely  abundant. 
Some  idea  of  their  numbers  can  be  obtained  when  it  is  known 
that  in  damp  places  or  along  the  banks  of  streams,  where  the 
butterflies  had  gathered  to  drink,  as  many  as  one  hundred  and 
fifty  individuals  were  counted  in  one  square  foot.  In  order  to 
estimate  the  numbers  flying  counts  were  made  of  the  individuals 
passing  between  two  fir  trees  about  twenty  feet  high  and  standing 
about  thirty  feet  apart.  The  counts  for  ten  successive  minutes 
between  4:40  and  4:50  P.M.  on  August  20,  1911,  were  as  follows: 


1st  minute  105 

2nd  minute  119 

3rd  minute  130 

4th  minute  102 

5th  minute  134 

6th  minute  ..  ..  100 


7th  minute  96 

8th  minute  102 

9th  minute  83 

10th  minute  112 

Av.  per  minute  ..  108 


It  can  readily  be  seen,  therefore,  that  butterflies  were  the 
insects  most  available  at  the  time.  The  significant  thing  is  that 
certain  birds  changed  their  food  habits  to  meet  the  changed  con- 
ditions. Birds  collected  in  the  same  locality  before  the  butter- 
flies became  abundant  had  taken  no  butterflies. 

The  investigation  did  not  show  that  birds  can  be  depended 
upon  to  control  butterfly  outbreaks,  for  the  numbers  taken  com- 
pared with  the  actual  numbers  of  the  insects  were  insignificant. 
The  fact  that  the  birds  attacked  the  insect  at  a  critical  time  in 
its  life-history — the  adult  stage  when  the  death  rate  is  at  its 
minimum  and  the  insect  has  the  best  chance  of  surviving  till  egg- 
laying — made  the  work  of  birds  more  important  than  if  they 


1914]   Bryant:  Economic  Status  of  the  Western  Meadowlark     459 

had  fed  upon  the  larvae  or  pupae.  However,  any  increased  de- 
struction at  the  time  when  insects  appear  in  abnormal  numbers 
must  be  considered  a  benefit. 

Four  out  of  the  five  birds  found  to  feed  on  the  butterflies 
are  numbered  among  the  birds  whose  usual  food  habits  justly 
subject  them  to  severe  criticism  from  the  farmer.  Consequently 
the  conclusion  can  be  drawn  that  some  of  the  birds  noted  for 
their  depredations  often  become  valuable  insect  destroyers  at  just 
the  time  when  they  are  most  needed  as  such. 

This  evidence  fails  to  support  Mr.  McAtee's  contention  that 
"butterflies  are  in  very  little  demand  with  birds  in  the  United 
States."  Nor,  on  the  other  hand,  does  it  support  the  conclusions 
of  Finn  (1897)  that  "there  is  a  general  appetite  for  butterflies 
among  insectivorous  birds,  even  though  they  are  rarely  seen  when 
wild  to  attack  them."  Middle  ground  seems  to  be  the  best 
position  to  assume  until  more  evidence  is  at  hand. 

In  our  attempt  to  explain  why  butterflies  are  seldom  taken 
by  birds  we  have  laid  emphasis  on  palatability.  This  factor  can 
at  best  be  but  one  of  many  factors,  many  of  which  are  perhaps 
just  as  important  if  not  more  important.  Then,  too,  it  should 
be  noted  that  palatability  is  an  extremely  variable  factor,  for 
it  varies  with  the  species,  the  time  of  year,  the  availability  of 
food,  etc.  We  still  need  a  full  examination  of  all  of  the  factors 
governing  the  food-taking  of  birds  in  order  to  explain  the  inter- 
relation between  birds  and  butterflies. 

In  an  investigation  of  a  grasshopper  outbreak  at  Los  Banos, 
Merced  County,  in  July,  1912,  it  was  found  that  western  meadow- 
larks  were  destroying  an  average  of  nearly  fifty  grasshoppers  a 
day.  From  point  of  numbers  and  average  number  of  grasshop- 
pers destroyed,  the  western  meadowlark  was,  next  to  the  bicolored 
red-wing,  the  most  efficient  destroyer  of  the  pests.  Meadowlarks 
averaged  more  grasshoppers  per  bird  and  were  outdone  by  the 
red-wings  only  when  the  numbers  of  individual  birds  and  the 
total  destruction  accomplished  by  each  species  were  considered. 

A  comparison  of  the  food  of  meadowlarks  taken  in  the  same 
locality  in  1911,  when  grasshoppers  were  not  so  abundant  as  they 
were  in  1912,  demonstrated  the  fact  that  meadowlarks  averaged 
more  grasshoppers  when  these  insects  were  abnormally  abundant 


460          University  of  California  Publications  in  Zoology    [VOL.  11 

than  they  did  when  they  were  not  so  abundant.     The  results  of 
stomach  examinations  follow : 


FOOD  OF  WESTERN  MEADOWLARK  AT  Los  BANGS,  MERCED  COUNTY, 
CALIFORNIA 


Number 
of  birds 

Date 

Anhm 
food 

10 

July 

11, 

22, 

1911 

99.0 

5 

July 

15, 

17, 

1912 

99.2 

Vegetable 
food 

1.0 


Average 
no.  of 
grass- 
hoppers 
per  bird 

7 
16 


Total 
per  cent 


hoppers 
83.1 
96.2 


Meadowlarks  took  very  nearly  the  same  percentage  (99,  99.2 
per  cent)  of  animal  food  each  year  at  the  same  season,  showing 
that  at  this  time  of  year  the  bird  is  almost  wholly  insectivorous. 
The  availability  of  grasshoppers  as  a  diet  appears  to  have  influ- 
enced the  birds  taken  in  1912,  for  they  averaged  sixteen  grass- 
hoppers apiece  as  against  seven  taken  by  birds  collected  in  1911. 

As  the  numbers  of  grasshoppers  in  1911,  compared  with  the 
numbers  in  1912,  is  not  definitely  known,  it  is  impossible  to  state 
whether  these  birds  changed  their  food  habits  in  response  to  the 
extreme  availability  of  the  insects  in  1912.  It  is  also  impossible 
to  state  whether  the  numbers  taken  in  1912  were  in  direct  pro- 
portion to  the  numbers  taken  in  1911  or  whether  they  failed  fully 
to  respond  to  the  change  in  insect  population.  The  fact  remains, 
however,  that  meadowlarks,  and  other  birds  as  well,  took  greater 
numbers  of  grasshoppers  when  they  were  abnormally  abundant, 
but  also  forsook  certain  articles  of  diet,  such  as  beetles  and  weed 
seeds,  thus  causing  an  increased  percentage  of  grasshoppers  to 
be  taken  as  food.  The  direction  of  the  change  of  food  habits 
was  certainly  coincident  with  the  direction  of  change  in  food 
supply. 

The  efficiency  of  a  bird  as  an  insect  destroyer  at  the  time  of 
an  insect  outbreak  is  governed  more  largely  by  the  numbers  of 
birds  than  by  their  individual  capacity.  This  was  conclusively 
shown  in  the  investigation  of  the  grasshopper  outbreak.  The 
comparative  destruction  of  grasshoppers  per  day  by  single  indi- 
viduals and  by  the  total  number  of  each  species  is  represented 
in  the  following  table: 


1914]   Bryant:  Economic  Status  of  the  Western  Meadowlark     461 


COMPARATIVE  DAILY  DESTRUCTION  OF  GRASSHOPPERS  BY  BIRDS 


Species 
Anthony  Green  Heron 

Kildeer 

Burrowing  Owl 
Western  Kingbird 
Black  Phoebe 
California  Horned  Lark 
Bicolored  Eed-wing 
Western  Meadowlark 
Bullock  Oriole 
Brewer  Blackbird 
English  Sparrow 
Cliff  Swallow 
California  Shrike 


Average 
hoppers 

no.  of  grass- 
per  day  by 

Class 
B 

One 
bird 

42 

Total 
population 

1,050 

B 

33 

5.445 

A 

84 

1,260 

D 

8 

1,280 

D 

9 

180 

:         D 

8 

88 

B 

29 

78,590 

B 

48 

24,720 

B 

45 

4,050 

D 

9 

225 

D 

2 

100 

D 

3 

2,265 

C 

12 

1,200 

;are  mile 

by  total 

120.453 

Relative 

destruction  by 

different  species 

represented 

by  lines 


Class  A  represents  birds  taking  an  average  of  over  50  grass- 
hoppers per  day;  class  B,  25-50;  class  C,  10-25;  and  class  D, 
fewer  than  10  grasshoppers. 

The  comparative  numbers  of  the  different  species  were  calcu- 
lated by  averaging  censuses  taken  and  by  using  the  average  per 
mile  as  a  multiplier.  Although  not  accurate,  the  table  never- 
theless demonstrates  the  fact  that  such  birds  as  the  bicolored 
red-wing  and  the  western  meadowlark,  birds  of  small  capacity, 
because  of  their  greater  numbers,  far  outrank  in  efficiency  birds 
with  larger  individual  capacity. 

The  fact  that  the  western  meadowlark,  or  any  bird,  turns 
its  attention  to  the  insect  most  abundant  only  emphasizes  its  value 
as  a  balancer.  If  meadowlarks  took  no  greater  proportion  of 
insects  when  they  are  in  abnormal  numbers  than  when  they  were 
in  normal  numbers,  they  would  play  a  decreasing  part  in  restoring 
a  balance.  Since  they  do  change  the  proportions  of  food  to  meet 
the  fluctuations  in  the  number  of  insects,  they  must  be  consid- 
ered an  important  factor  in  the  restoration  of  normal  conditions. 


462          University  of  California  Publications  in  Zoology    [VOL.  n 

A  great  number  of  factors  operating  together  determine  the 
abundance  of  an  insect.  Birds  and  other  natural  enemies  are 
but  one  of  these  many  factors.  The  rate  of  reproduction  and 
food  supply  are  probably  more  important  factors.  In  spite  of 
this  fact  birds  are  one  of  the  limiting  factors  and  are  deserving 
of  attention  as  such. 

The  following  facts  have  been  demonstrated  by  these  investi- 
gations : 

1.  Birds  cannot  be  considered  a  dependable  means  of  com- 
pletely controlling  all  insect  outbreaks,  but  can  be  inferred  to  be 
instrumental  in  the  prevention  of  many. 

2.  Birds  can  be  depended  upon  to  act  as  defenders  and  pro- 
tectors of  crops  because  of  their  warfare  against  insect  pests. 

3.  Birds  change  their  food  habits  and  feed  on  the  insect  most 
abundant,  thereby  making  themselves  important  maintainers  of 
the  desired  balance  in  nature. 

4.  The  failure  of  birds  to  check  an  insect  outbreak  entirely 
is  evident  to  all.     Their  success  in  preventing  insects  from  be- 
coming abundant  is  not  so  apparent,  but  is  none  the  less  real. 
All  obtainable  evidence  points  to  the  fact  that  the  regulative 
influence  exerted  by  birds  when  insects  are  to  be  found  in  normal 
numbers,  though  less  apparent,  is  none  the  less  important,  for 
at  such  times  artificial  control  measures  are  seldom  used. 

5.  Birds,  which  on  account  of  their  abundance  cause  serious 
losses  to  the  agriculturist,  often  become  for  the  same  reason  the 
most  efficient  insect  destroyers  at  the  time  of  an  insect  outbreak. 

6.  Birds  help  to  maintain  an  equilibrium  in  nature.     Their 
destruction,  therefore,  causes  a  dangerous  disturbance  of  that 
balance  of  nature  most  suited  to  mankind. 

VERDICT  OF  RANCHERS 

In  order  that  the  opinion  of  the  men  most  directly  concerned 
might  not  be  overlooked,  a  circular  letter  was  sent  out  to  promi- 
nent ranchers  throughout  the  state.  A  copy  of  the  letter  follows : 

DEAR  SIR: — 

The  State  Fish  and  Game  Commission  has  taken  up  the  study  of  the 
meadowlark  in  its  relation  to  agriculture  and  desires  to  know  what  you 
think  of  the  bird.  In  order  to  secure  comprehensive  and  uniform  data, 
answers  to  the  following  questions  are  urgently  requested: 


1914]   Bryant:  Economic  Status  of  the  Western  Meadoivlark     463 

1.  Name  

2.  Address  

3.  Occupation  

4.  How  many  acres  of  land  do  you  own?  

Is  your  ranch  hilly  upland  or  bottom  land?  

5.  What  is  the  principal  crop  raised?  

What  other  crops?  

6.  Has  the  meadowlark  done  any  harm  on  your  place?  

If  so,  how  and  to  what  extent?  

7.  Have  you  examined  the  stomachs  of  any  meadowlarks  to  ascertain  their 

food?  If  so,  what  was  in  the  stomachs?  

8.  Approximately,  how  many  meadowlarks  are  seen  daily  on  your  place? 

Are  the  numbers  any  greater  when  the  grain 

is  sprouting?  

9.  Do  you  prize  the  meadowlark  as  a  song  bird?  

10.  On  the  whole,  do  you  consider  the  meadowlark  a  nuisance?  

Any  additional  information  that  you  can  give  on  the  subject  will  be 
appreciated.  Address  all  communications  to  H.  C.  Bryant,  Assistant  State 
Fish  and  Game  Commission,  East  Hall,  University  of  California,  Berkeley, 
California. 

Over  a  hundred  replies  to  this  letter  were  received.  Although 
the  returns  may  be  criticised  on  the  grounds  that  a  greater 
number  of  those  interested  in  the  bird  because  of  its  esthetic 
value  sent  in  answers,  yet  care  was  taken  to  avoid  this.  Blanks 
were  sent  to  the  men  who  complained  of  the  depredations  of  the 
meadowlark  and  to  ranchers  irrespective  of  their  particular  point 
of  view.  The  average  acreage  of  the  men  reporting  was  638,  so 
that  it  can  be  seen  that  the  verdict  is  not  from  small  land-holders 
or  orchardists.  Over  ninety-eight  per  cent  of  those  reporting 
grew  grain  or  hay. 

When  the  returns  as  to  whether  the  meadowlark  is  a  nuisance 
and  as  to  whether  it  damages  crops  are  classified  as  to  counties 
the  results  are  as  follows : 

Is  the  meadowlark  Does  the  meadowlark 

a  nuisance?  damage  crops? 

Number  , A ^  , * N 

County                   reporting               Yes                No  Yes                 No 

Siskiyou  5                  5                  5 

Shasta  1                 1                 1 

Humboldt  2                 2                 2 

Trinity  5  1?  4  5 

Mendocino  1  1              1              

Tehama  3  21  21 

Sutter  55  5 


464          University  of  California  Publications  in  Zoology    [VOL. 11 


Number 
County                   reporting 

Plumas                             1 

Is  the  meadowlark 
a  nuisance  ? 

Does  the  meadowlark 
damage  crops? 

'Yes 

No    ^ 
1 

Yes 

No   ^ 
1 

Butte 

2 

2 



2 

Yolo 

1 

1 



1 

Sonoma 

4 

4 



4 



Napa 

2 

1 

1 

1 

1 

Sacramento 

2 

1 

1 

1 

1 

Marin 

1 

1 

1 



Contra  Costa 

1 

1 

1 

Alameda 

7 

2 

4 

3 

4 

San  Joaquin 

17 

11 

5 

11 

6 

Stanislaus 

2 



2 

2 

El  Dorado 

1 

1 



1 

Calaveras 

1 

1 



1 



Madera 

1 

1 



1 

Merced 

3 

2 

1 

2 

Fresno 

5 

4 

1 

4 

1 

Monterey 

9 

2 

7 

2 

6 

San  Luis  Obispo 

1 

1 



1 

Kings 

5 

3 

1 

4 

1 

Tulare 

4 

1 



3 

1 

Mono 

3 



3 



3 

Inyo 

9 

2 

7 

2 

7 

Kern 

1 



1 

Santa  Barbara 

3 

3 

3 

Ventura 







Los  Angeles 

2 

2 

2 

San  Bernardino 

1 

1 

1 

Orange 

5 



4 

5 

Eiverside 

4 

1 

3 

2 

2 

San  Diego 

2 



2 

0 

Imperial 





Totals 

122 

48 

65 

54 

67 

Total  coast 
counties 

31 

5 

25 

5 

25 

Total  centra] 
counties 

60 

35 

19 

38 

22 

Total  northern 
California 

105 

47 

50 

52 

52 

Total  southern 
California 

17 

1 

15 

2 

15 

1914]   Bryant:  Economic  Status  of  the  Western  Meadowlark     465 

It  will  be  seen  from  this  table  that  there  is  a  considerable 
difference  of  opinion  as  to  whether  the  western  meadowlark 
damages  crops.  The  astonishing  fact  is  that  many  grain  growers 
in  the  Sacramento  and  San  Joaquin  valleys  report  that  western 
meadowlarks  do  not  injure  their  crops,  or  that  the  injury  is 
negligible. 

The  kinds  of  crops  reported  as  being  damaged  ranged  from 
garden  truck,  melons  and  grapes  to  corn  and  sprouting  grain. 
Damage  to  garden  truck,  melons  and  grapes  was  reported  by 
three  or  fewer  men.  Damage  to  oats  was  reported  by  over 
twenty,  barley  by  less  than  this  number,  and  wheat  by  less  than 
ten.  Most  of  the  reports  did  not  designate  the  kind  of  grain, 
simply  stating  that  meadowlarks  damaged  sprouting  grain.  An- 
swers as  to  the  extent  of  damage  varied  from  "none"  to  "total 
crop."  The  number  of  meadowlarks  seen  was  reported  as  being 
from  two  or  three  up  into  the  thousands.  The  answers  to  this 
question  cannot  be  considered  reliable.  A  large  majority  of  those 
who  considered  the  meadowlark  a  nuisance  answered  the  question 
whether  the  meadowlark  was  prized  as  a  song  bird  in  the  nega- 
tive, whereas  those  answering  the  former  question  in  the  negative 
almost  unanimously  answered  the  latter  in  the  affirmative. 

Reports  of  damage  were  most  numerous  from  the  Sacramento 
and  San  Joaquin  valleys.  This  seems  natural,  for  grain  is  the 
crop  most  widely  grown  in  this  section  and  meadowlarks  are 
most  numerous.  Southern  California  is  most  unanimous  in  its 
verdict  of  "not  guilty."  Two  reasons  can  be  made  to  account 
for  this:  the  comparatively  small  amount  of  grain  raised  and 
the  comparatively  small  number  of  meadowlarks.  Few  reports 
of  damage  have  come  from  the  northern  coast  region,  in  spite 
of  the  fact  that  meadowlarks  are  very  numerous  in  this  section. 

The  majority  of  those  reporting  have  not  had  crops  damaged 
by  meadowlarks  and  do  not  consider  the  bird  a  nuisance.  It  does 
not  seem  reasonable  to  believe  that  all  of  these  men  based  their 
report  on  sentiment.  Evidence  seems  to  point  rather  to  the  fact 
that  many  of  those  complaining  of  damage  have  based  their  judg- 
ment on  circumstantial  evidence  and  have  somewhat  exaggerated 
the  real  damage  done. 


466  University  of  California  Publications  in  Zoology    [VOL.  11 

A  DETERMINATION  OF  THE  ECONOMIC  STATUS  OF  THE  WESTERN 
MEADOWLARK  IN  CALIFORNIA 

Field  investigation  of  the  damage  to  crops  has  led  to  the 
following  conclusions: 

1.  The  western  meadowlark  is  destructive  to  sprouting  grain 
because  of  its  habit  of  boring  down  beside  the  sprout  and  pulling 
off  the  kernel.    The  amount  of  damage  varies  with  the  location, 
the  abundance  of  the  birds,  the  time  of  year,  the  character  of 
the  soil,  and  the  kind  of  grain.    The  damage  to  oats  is  greatest ; 
wheat  suffers  less  and  barley  little.     A  greater  loss  can  be  ex- 
pected with  broadcasted  grain  than  with  drilled  grain,  because 
not  being  sowed  so  deeply  it  is  more  readily  obtained  by  meadow- 
larks.     The  real  amount  of  damage  done  has   evidently  been 
overestimated,  however,  for  fields  apparently  badly  damaged  have 
given  the  average  yield  later  in  the  year.     On  the  other  hand, 
where  meadowlarks  are  very  numerous  and  the  quantity  of  grain 
small,  fields  have  had  to  be  resown  to  assure  a  crop. 

2.  In  the  destruction  of  sprouting  grain  we  have  the  only 
serious  count  against  the  meadowlark,   for  damage  to  melons, 
grapes   and  other  crops  has  been  found  to  be  negligible.     A 
number  of  things  minimize  the  damage  done,  chief  of  which  is 
the  fact  that  meadowlarks  are  able  to  obtain  the  kernel  for  a 
limited  period  only.     After  the  second  and  third  leaves  have 
appeared,  the  plant  is  well  rooted  and  the  loss  of  the  kernel  does 
not  destroy  the  plant.     Hence  damage  is  limited  to  a  period  of 
about  two  weeks  on  any  given  field  and  is  reduced  by  deep 
planting. 

3.  Those  factors  which  make  the  depredations  of  the  western 
meadowlark  important   and   those   factors   which   minimize   the 
damage  done  may  be  summarized  as  follows : 

1.  Method    of    pulling    sprouting  1.  Same  method  valuable  in  se- 

grain.  curing  such  insects  as  cut- 

2.  Lack  of  insect  food  when  grain  worms  and  wireworms. 

is  sprouting  coupled  with  the  2.  Take   a   larger   percentage   of 

availability  of  grain  at  the  insects  than  of  grain  during 

same  time.  year. 

3.  Flocking  habit.  3.  Apparently    driven    to    grain 

only   when   insects   are    not 
available. 


1914]   Bryant:. Economic  Status  of  the  Western  Meadowlark     467 


4.  Abundance  of  meadowlarks  in 

grain-growing  localities. 

5.  Great  capacity  and  rapid  diges- 

tion. 


4.  Time  during  which  damage  can 

result  limited. 

5.  Flocking  habit  makes  control 

measures  easier. 

6.  Abundance  of  meadowlarks  as- 

sures more  efficient  destruc- 
tion of  insect  pests. 

7.  Great  capacity  and  rapid  diges- 

tion improves  their  value 
as  insect  destroyers;  slower 
digestion  of  grain  than  of 
insects  makes  a  less  con- 
sumption of  the  former. 

8.  Do  not  destroy  other  crops. 

9.  A  certain  amount  of  thinning 

is  sometimes   desirable. 

10.  Often  perform  service  in   de- 

stroying insects  in  same  field 
where  damage  was  done. 

11.  Prefer   uncultivated   to    culti- 

vated land. 

12.  Unable  to  cause  serious  damage 

when  grain  is  planted  deeply. 


4.  A  study  of  the  life-history  of  the  western  meadowlark 
shows  it  to  be  a  bird  which  prefers  uncultivated  to  cultivated 
land,  especially  while  feeding  and  nesting.     It  feeds  in  places 
where  other  birds  do  not  feed,  and  takes  many  of  the  ground- 
loving  insects  which  other  birds  do  not  take.    Its  habit  of  boring 
into  the  ground  after  food  makes  it  important  as  a  destroyer 
of  such  insects  as  cutworms,  wireworms,  and  tipulid  larvae,  very 
destructive  insects  of  the  grain  fields  and  meadows.    Young  birds 
are  fed  entirely  on  insects  and  need  nearly  their  own  weight  of 
food  each  day.     They  demand  the  largest  amount  of  food  when 
insects  are  at  a  maximum.     The  western  meadowlark  appears 
to  increase  in  numbers  with  cultivation  of  land  when  a  proper 
food  supply  is  furnished.     It  can  be  seen,  therefore,  that  most 
of  the  facts  regarding  its  life-history  tend  to  place  it  among  the 
beneficial  birds. 

5.  Experimentation  has  shown  that  western  meadowlarks,  like 
other  birds,  have  a  very  rapid  digestion  and  are  able  to  digest 
a  full  meal  in  four  to  six  hours.    Under  these  circumstances  it 
is  evident  that  they  must  consume  large  quantities  of  food.    Ex- 


468          University  of  California  Publications  in  Zoology    [VOL. 11 

perimental  feeding  has  substantiated  this  fact.  The  larger  the 
amount  of  food  consumed  the  larger  must  be  the  toll  taken.  As 
experiment  showed  that  insects  are  digested  more  rapidly  than 
grain,  proportionately  larger  amounts  of  insect  food  than  vege- 
table food  may  be  consumed  daily. 

6.  Stomach  examination  has  demonstrated  the  following  facts : 
(1)  A  much  larger  percentage  of  animal  food  (sixty  per  cent) 
is  taken  during  the  year  than  vegetable  food  (forty  per  cent). 
The  meadowlark  feeds  on  grain  to  a  considerable  extent  during 
the  winter  months.  A  very  small  proportion  of  that  found  in 
the  stomachs  was  sprouted  grain.  A  large  proportion  of  the 
grain  taken  was  made  up  of  wild  oats  and  had  evidently  not 
been  gathered  in  grain  fields.  If  meadowlarks  by  their  depre- 
dations can  drive  many  of  the  ranchers  of  the  state  to  drill  their 
grain  and  plant  it  more  deeply  they  will  be  performing  a  service 
by  increasing  the  yield.  The  increased  yield  so  produced  would 
doubtless  more  than  cover  the  loss  sustained  from  the  birds. 

(2)  Western  meadowlarks  destroy  large  numbers  of  insects 
which   are   injurious  to  the  same  crop   damaged   by  the  birds 
themselves. 

(3)  The  kind  of  food  varies  with  time  of  year,  locality,  and 
abundance  of  available  food.     This  correlation  of  food  habits 
with  the  environmental  conditions  increases  the  bird's  value  as 
a  balancer.     If  the  meadowlark  did  not  change  its  food  with  a 
change  in  supply,  its  value  as  an  insect  destroyer  would  be  slight. 
In  turning  its  attention  to  the  food  most  abundant  it  becomes 
an  important  factor  in  maintaining  a  balance.     Its  survival  as 
a  destroyer  of  insects  is  dependent  on  its  being  able  to  obtain  a 
supply  of  food  when  animal  food  is  not  available. 

(4)  The  quantity  of  injurious  insects  taken  daily  is  large 
enough  to  make  this  bird  of  at  least  some  importance  as  a  de- 
stroyer of  insect  pests.    The  cumulative  effect  of  such  destruction 
enhances  the  bird's  value.    Few  beneficial  insects  are  destroyed. 

(5)  The  meadowlark  destroys  quantities  of  seeds  of  serious 
weed  pests. 

The  service  which  birds  render  to  agriculture  has  been  over- 
emphasized by  those  appreciating  the  esthetic  value  of  birds. 


1914 j   Bryant:  Economic  Status  of  the  Western  Meadowlark     469 

Such  writers  have  been  prone  to  take  the  point  of  view  that 
birds  were  expressly  made  to  destroy  injurious  insects.  One 
might  just  as  well  say  that  insects  were  created  to  furnish  food 
for  birds.  If  birds  should  become  numerous  enough  actually  to 
control  the  number  of  insects,  they  would  doubtless  become  a 
greater  pest  than  the  insects  themselves. 

Parasitic  insects  are  not  proving  to  be  the  controls  which 
their  advocates  have  maintained  they  would  be.  And  certainly 
there  are  arguments  which  prove  that  birds  are  not  the  panacea 
for  all  insect  ills.  Birds  destroy  many  of  the  most  beneficial 
insects  known.  Some  of  the  most  injurious  insects  are  less  often 
taken  by  birds.  Birds,  though  they  be  in  abundance,  fail  to 
prevent  outbreaks  of  injurious  insects.  Birds  scatter  weed  seed 
as  well  as  destroy  it.  If  there  were  no  birds,  would  not  other 
factors,  such  as  parasitism,  climatic  conditions,  etc.,  soon  bring 
about  a  balance?  Besides,  cannot  insect  pests  be  more  surely 
and  successfully  controlled  by  artificial  means,  insecticides, 
sprays,  etc.  ?  On  the  other  hand,  is  there  not  a  saving  of  expense 
in  letting  nature  control  insect  ravages  as  far  as  possible  ? 

All  of  these  points  deserve  our  consideration.  They  need  to 
be  weighed  in  the  balance.  When  all  the  evidence  for  and 
against  the  utility  of  birds  is  in,  a  solution  will  be  available. 
Until  that  time  there  will  always  be  two  sides  to  the  question. 

A  partial  solution  of  the  problem  is  afforded  by  placing 
emphasis  elsewhere,  thereby  avoiding  these  two  opposing  sides 
of  the  question.  Forbes  (1903)  pointed  out  that  the  value  of 
birds  does  not  lie  in  the  fact  that  they  discriminate  and  take 
only  injurious  insects,  but  in  the  fact  that  they  eat  insects.  The 
place  filled  by  birds  in  the  economy  of  nature  is  the  important 
thing. 

Most  life  under  the  natural  order  of  things  is  conditioned 
very  largely  by  its  food  supply:  in  the  case  of  purely  insectiv- 
orous birds,  by  insects;  in  the  case  of  insects,  by  plant  life.  If 
it  be  true,  as  it  appears  to  be,  that  organisms  become  so  adjusted 
to  their  food  supplies  that  only  the  surplus  or  excess  is  normally 
taken,  then  the  importance  of  birds  in  their  relation  with  insects 
lies  in  their  toll  of  the  surplus.  Since  it  is  the  excess  or  abnormal 
abundance  of  insects  that  makes  most  trouble  for  the  agricul- 


470          University  of  California  Publications  in  Zoology    [VOL.  11 

turist  and  horticulturist,  any  factor,  even  though  it  be  one  of 
many  factors,  is  important  in  insect  control.  Birds  destroy 
insects;  they  are  therefore  a  natural  factor  in  the  control  of 
insect  life.  Since  they  do  not  fluctuate  in  numbers  so  greatly 
as  do  parasitic  insects,  they  are  a  more  constant  factor  than 
parasites.  Insectivorous  birds  feed  on  the  insect  most  abundant, 
hence  they  are  the  more  important  in  limiting  the  numbers  of 
insects.  The  proportion  taken  apparently  varies  directly  as  the 
numbers  of  the  insects.  The  greatest  toll  on  insect  life  comes 
during  the  nesting  season,  a  time  when  the  insect  population  is 
at  its  maximum.  This  in  itself  establishes  the  fact  that  birds 
have  a  real  and  an  important  part  to  play  in  the  interaction  of 
organisms. 

Almost  all  insects  are  potentially  injurious.  Injurious  insects 
in  small  numbers  cause  practically  no  damage.  Neutral  and 
beneficial  insects  in  large  numbers  may  become  injurious.  Hence 
the  destruction  of  neutral  and  beneficial  insects,  if  they  are 
potentially  destructive,  may  become  at  times  of  utility. 

Insects  have  in  turn  adapted  themselves  to  the  constant  drain 
on  their  numbers.  This  becomes  very  evident  when  we  study 
the  rate  of  reproduction  of  insects.  However,  it  is  the  balance 
we  need.  Birds  and  insects  both  have  a  part  to  play  in  the 
balance.  They  are  supplementary  and  indispensable. 

This  is  an  important  viewpoint.  Emphasis  on  the  relation 
of  birds  to  insects  viewed  from  the  standpoint  of  the  interaction 
of  organisms  is  rightly  placed.  Birds  are  important  because  they 
evidently  are  an  indispensable  factor  in  maintaining  an  equi- 
librium of  organic  life.  If  all  birds  play  this  important  part, 
the  destruction  of  any  particular  species  of  bird  means  a  dis- 
turbance in  the  balance.  The  nearer  man  adjusts  organic  life 
to  his  desires  the  more  important  will  become  each  natural  factor 
concerned.  Artificial  means  of  adjustment  often  lack  the  effi- 
ciency of  natural  means. 

It  is  readily  acknowledged  that  birds  are  not  the  only  checks 
on  the  increase  of  insects.  The  very  large  toll  taken  by  them, 
however,  places  them  in  the  front  rank  as  insect  destroyers. 
Parasites  can  become  abundant  only  when  their  host  becomes 
abundant  and  do  their  work  effectively  only  after  the  insect  has 


1914]   Bryant:  Economic  Status  of  the  Western  Meadowlark     471 

had  sufficient  time  to  cause  damage.  Birds  in  order  to  keep  alive 
must  wage  a  continual  warfare  on  insect  life,  no  matter  what 
the  abundance.  They  are  evidently,  therefore,  to  be  relied  upon 
as  more  dependable  regulators  than  parasites. 

Since,  as  has  been  shown,  the  average  adult  western  meadow- 
lark  destroys  nearly  three  pounds  of  insects  each  year  and  prob- 
ably almost  as  many  more  pounds  while  feeding  its  young,  its 
value  to  the  agriculturist  is  apparent.  The  ratio  of  value  of  one 
of  these  birds  living  to  that  of  one  dead  is,  therefore,  as  five 
pounds  of  insects  and  one-half  pound  of  weed  seeds  are  to  one 
and  three-fourths  pounds  of  grain,  a  considerable  portion  of 
which  is  made  up  of  wild  oats  and  waste  grain. 

The  fact  that  the  western  meadowlark  destroys  certain  bene- 
ficial insects  cannot  be  counted  a  point  in  its  favor.  And  yet 
the  quantity  taken  is  so  small,  less  than  five  per  cent  of  the  food 
for  the  year,  and  the  destruction  so  caused  is  such  an  indirect 
injury,  that  the  damage  possible  is  very  slight  and  practically 
negligible.  The  destruction  of  one  ichneumon-fly  compared  with 
the  destruction  of  one  hundred  grasshoppers,  somewhat  the  pro- 
portion in  which  they  are  taken,  leaves  no  doubt  as  to  the  com- 
parative benefit. 

Since  the  western  meadowlark  feeds  to  a  large  extent  on 
grass-land  insects,  many  of  which  are  not  eaten  by  other  birds, 
it  must  be  considered  a  friend  of  the  dairymen  and  grain  growers, 
and  not  an  enemy.  "The  laborer  is  worthy  of  his  hire."  The 
grain  taken  by  western  meadowlarks  can  well  be  considered  the 
pay  for  their  efficient  work  in  destroying  injurious  insects  and 
weed  seed. 

It  must  be  apparent  from  these  comparisons  that  the  balance 
is  certainly  in  favor  of  the  meadowlark.  Birds  are  considered 
a  national  resource  and  so  belong  to  the  people  as  a  whole.  It 
seems  doubtful  whether  the  grain  grower  should  destroy  birds 
destroying  his  crops  when  the  same  birds  might  be  performing 
a  great  service  in  destroying  injurious  insects  in  his  own  or  his 
neighbor's  alfalfa  field. 

This  investigation  has  shown  that  the  western  meadowlark, 
as  a  rule,  deserves  protection  and  encouragement  at  the  hands 
of  the  agriculturist.  Only  in  rare  cases  can  it  be  said  that  the 


472          University  of  California  Publications  in  Zoology    [VOL.  11 

bird  does  more  harm  than  good.  One  and  three-fourths  quarts 
of  insects  taken  by  a  western  meadowlark  during  a  year  more 
than  pay  for  less  than  one  quart  of  grain,  a  large  part  of  which 
does  not  represent  a  loss. 

Although  emphasis  has  been  laid  on  the  dollar  and  cents 
value,  yet  the  other  values  which  cannot  be  reckoned  on  the 
money  basis  must  be  taken  into  account.  And  here  we  find  an 
even  stronger  defense  of  the  western  meadowlark,  for  the  esthetic 
and  scientific  values  greatly  strengthen  the  case  for  the  bird,  in 
spite  of  the  fact  that  these  values  are  often  unacknowledged  by 
many  who  profit  by  them. 

The  investigation  has  shown  that  for  the  following  ten  reasons 
the  western  meadowlark  should  remain  a  protected  non-game 
bird. 


TEN   REASONS   WHY   THE   WESTERN   MEADOWLARK 

(STURNELLA  NEGLECT A)   SHOULD  BE  A 

PROTECTED  NON-GAME  BIRD 

1.  As  a  destroyer  of  cutworms,  caterpillars,  and  grasshoppers, 
three  of  the  worst  insect  pests  in  the  State  of  California,  the 
western  meadowlark  is  probably  unequaled  by  any  other  bird. 
The  stomachs  of  meadowlarks  examined  have  averaged  as  high 
as  six  cutworms  and  caterpillars,  and  sixteen  grasshoppers  apiece. 
Maximum  numbers  of  sixty-six  cutworms  and  of  thirty-two 
grasshoppers  have  been  taken  from  a  single  stomach.  As  the 
time  of  digestion  is  about  four  hours,  three  times  the  average 
must  be  consumed  daily.  Other  injurious  insects  destroyed  are 
click-beetles,  the  larvae  of  which  are  wireworms,  May  beetles, 
weevils,  crickets,  Jerusalem  crickets,  stink-bugs,  flies,  and  crane- 
flies.  Fifty-nine  and  six-tenths  per  cent  of  the  food  for  the  year 
is  made  up  of  animal  food.  Each  meadowlark  in  the  state  con- 
sumes, at  the  least  calculation,  six  pounds  of  food  each  year,  two 
and  three-fourths  pounds  of  which  is  made  up  of  insects,  most 
of  which  are  injurious  to  crops. 


1914]   Bryant:  Economic  Status  of  the  Western  Meadowlark     473 

2.  The  western  meadowlark  destroys  a  very  small  percentage 
of  beneficial  insects.     That  one  hundred  grasshoppers  are  de- 
stroyed to  every  parasitic  ichneumon-fly  is  a  very  conservative 
estimate. 

3.  But  one  crop  is  attacked,  for  the  only  serious  loss  occa- 
sioned by  the  western  meadowlark  is  to  sprouting  grain.    Damage 
to  fields  of  sprouting  grain  can  be  largely  prevented  by  certain 
protective  measures,  some  of  which  have  been  proved  by  experi- 
ment to  be  instrumental  in  producing  better  crops.    Hence  injury 
to  crops  is  not  only  limited  but  largely  preventable. 

4.  The  western  meadowlark  destroys  the  seeds  of  some  of 
California 's  worst  weed-seed  pests.    During  the  fall  months,  when 
insects  are  not  available,  this  bird  destroys  large  quanties  of 
weed  seeds.    Over  two  hundred  weed  seeds  have  been  taken  from 
a  single  stomach.    Napa  thistle,  Johnson  grass,  canary  grass,  fox- 
tail, tarweed,   pigweed,  tumbleweed,   mustard,   turkey  mullein, 
sunflower,  and  nightshade  are  among  the  weed  pests  destroyed. 

5.  The  western  meadowlark,  by  feeding  in  places  not  fre- 
quented by  other  birds,  procures  many  injurious  insects  which 
would  not  fall  prey  to  other  animals.    In  so  doing  it  fills  a  niche 
in  the  economy  of  nature  which  apparently  is  not  filled  by  any 
other  form  of  life. 

6.  Investigations  of  the  relations  of  birds  to  insect  outbreaks 
have  demonstrated  that  the  western  meadowlark,  by  turning  its 
attention  to  the  insect  most  available,  becomes  important  as  a 
maintainer  of  a  balance  of  insect  life.     As  such  it  becomes  a 
defender  and  protector  of  crops. 

7.  The  western  meadowlark  has  great  esthetic  value.    A  bird 
of  the  meadow  and  pasture,  it  adds  life  and  interest  to  treeless 
areas.    Its  bright  colors  and  beautiful  song  have  made  it  one  of 
the  best  known  birds  in  the  state.     The  meadowlark  even  adds 
to  the  value  of  suburban  real  estate. 

8.  The  western  meadowlark  cannot  be  considered  a  game  bird. 
The  following  quotation  is  from  an  article  by  D.   G.   Elliott 
(1864),  entitled  "The  Game  Birds  of  the  United  States" :    "Not 
indeed  every  feathered  biped,  which  a  high  breed  of  dogs  will 
instinctively  point,  can  be  included  in  our  list ;  for  the  meadow- 
lark    (Sturnella  magna),  that  troublesome  pest  of  every  true 


474          University  of  California  Publications  in  Zoology    [VOL.  11 

sportsman,  whose  dog,  unless  taught  otherwise,  will  surely  follow, 
has  fairly  no  claim  to  this  title,  any  more  than  have  a  snake 
or  a  turtle,  to  either  of  which  a  point  will  generally  be  made, 
and  these  last,  it  is  hardly  necessary  to  add.  are  neither  birds 
nor  game. "  In  some  of  the  eastern  states,  where  the  meadowlark 
is  still  unprotected,  a  parasitic  worm  often  found  in  the  small 
of  the  back  deters  many  experienced  persons  from  using  the 
bird  for  food.  The  western  meadowlark  Cannot  be  considered  a 
game  bird  under  the  definition  of  such  birds  given  in  Section 
637a  of  the  Penal  Code  of  the  State  of  California. 

9.  By  a  decision  of  the  United  States  Supreme  Court  (G-eer 
v.  Conn:,  161  U.  S.  519),  birds  are  considered  a  national  resource 
belonging  to  the  people   as   a   whole.     The   destruction  of  the 
western  meadowlark  by  the  grain  grower  causes  a  loss  to  the 
growers  of  alfalfa  and  other  crops  who  profit  enormously  by  the 
destruction  of  insect  pests  by  the  bird. 

10.  In  certain  sections  of  California,  notably  southern  Cali- 
fornia, because  of  the  small  numbers  of  meadowlarks  or  owing 
to  the  kind  of  crops  raised,  the  western  meadowlark  causes  no 
damage  and  is  considered  distinctly  beneficial.     The  extent  of 
damage  varies  with  the  locality  and  the  kind  of  crop  raised. 


SUGGESTIONS  FOR  THE  PROTECTION  OF  CROPS 

In  a  study  of  this  kind  something  should  be  said  as  to  the 
methods  of  protecting  crops  from  the  depredations  of  the  western 
meadowlark,  for,  although  we  may  in  general  say  that  the  bird 
is  highly  beneficial  to  agricultural  interests,  yet  local  conditions 
and  an  overabundance  of  birds  may  demand  protective  measures. 
Scarecrows  have  proved  inefficient  as  a  means  of  protecting  grain 
fields  from  attack  by  meadowlarks.  The  birds  soon  become  accus- 
tomed to  any  object  placed  in  the  field,  and  so  continue  their 
depredations.  Frightening  the  birds  by  shooting  is  found  to 
be  a  better  means,  but  is  not  always  practical.  It  should  be 
remembered  that  the  protective  measures  suggested  are  all  prac- 
tical ones,  for  it  has  been  shown  that  the  damage  is  limited  to 
two  weeks ;  also  that  a  certain  amount  of  thinning  is  allowable, 
and  in  some  cases  of  value.  The  outlav  of  several  extra  sacks 


1914]   Bryant:  Economic  Status  of  the  Western  MeadowlarJc     475 

of  grain  can  be  more  than  counter-balanced  by  the  efficiency  of 
the  meadowlarks  later  in  the  year  when  injurious  insects  have 
become  abundant  in  grain  fields. 

Where  losses  to  crops  warrant  protective  measures  the  fol- 
lowing are  proposed : 

1.  Plant  grain  deeply.    It  assures  a  better  crop  regardless  of 
losses  due  to  meadowlarks.     Drilled  grain  gives  a  better  yield 
than  broadcasted  and  is  also  better  protected  from  the  attack  of 
meadowlarks. 

2.  Fields  bordering  pasture  or  uncultivated  land,  if  sowed 
more  heavily   along  such  margins,   will  usually  be   assured  a 
normal  crop. 

3.  Meadowlarks  are  easily  frightened  from  a  field  by  the 
noise  of  shooting  or  by  a  dog.    As  damage  is  limited  to  a  short 
period  of  time,  this  method  seems  practical  on  small  fields. 

EECOMMENDATIONS  AS  TO  LEGISLATION 

The  United  States  Supreme  Court  (Geer  v.  Conn.,  161  U.  S. 
519)  has  ruled  that  all  game  and  wild  birds  belong  to  the  people. 
The  decision  of  the  court  in  a  test  case  in  California  is  as  follows : 
"We  take  it  to  be  the  correct  doctrine  in  this  country  that  the 
ownership  of  wild  animals,  so  far  as  they  are  capable  of  owner- 
ship, is  in  the  State,  not  as  a  proprietor,  but  in  its  sovereign 
capacity  as  the  representative  and  for  the  benefit  of  all  its  people 
in  common  (State  v.  Rodman,  /.  c.).  Consequently  the  people, 
through  their  various  legislatures,  control  this  natural  resource. 
It  is  evident,  therefore,  that  as  in  other  things,  the  majority  rule. 
In  legislation  the  rights  of  the  many  are  often  clouded  over  by 
the  activities  of  the  few.  Sometimes  this  is  due  to  inactivity 
on  the  part  of  the  many ;  sometimes  it  is  due  to  a  lack  of  knowl- 
edge on  the  part  of  the  many  and  an  accurate  knowledge  by  the 
few.  In  the  case  before  us  we  desire  that  there  shall  be  no  lack 
of  information  on  either  side. 

The  desire  on  the  part  of  certain  ranchers  of  the  state  to 
place  the  western  meadowlark  on  the  unprotected  list  in  the 
hope  that  it  might  be  so  reduced  in  numbers  as  to  prevent  injury 
to  crops,  and  the  desire  of  certain  sportsmen  to  add  this  bird 


476          University  of  California  Publications  in  Zoology    [VoL-  n 

to  the  list  of  game  birds,  has  twice  occasioned  the  introduction 
of  a  bill  into  the  State  Legislature.  The  same  influences  caused 
the  Fish  and  Game  Protective  Association  in  1912  to  recommend 
placing  this  bird  on  the  game  list.  In  spite  of  the  failure  of 
the  former  bills  to  pass,  protest  has  continued.  As  a  final  solution 
of  the  problem  this  investigation  was  ordered  by  the  State  Fish 
and  Game  Commission. 

*********** 

This  investigation  having  been  completed  and  dependable  data 
being  at  hand,  the  following  considerations  appear  to  the  writer 
to  demand  the  accompanying  recommendation  for  legislation : 

The  western  meadowlark  (Sturnella  neglecta),  along  with 
other  birds,  must  be  considered  an  important  resource  of  this 
state,  and  therefore  cannot  receive  destruction  at  the  hands  of 
some  without  distinct  loss  to  others. 

The  western  meadowlark  can  in  no  way  be  considered  a 
desirable  game  bird,  but  must  be  numbered  with  the  insectivor- 
ous non-game  birds. 

The  western  meadowlark,  through  a  thorough  scientific  inves- 
tigation lasting  over  a  period  of  over  two  years  and  including 
field  study  and  an  examination  of  nearly  two  thousand  stomachs, 
some  being  collected  each  month  of  the  year  in  over  twenty 
different  localities  in  California,  has  been  shown  to  be  distinctly 
beneficial  to  agricultural  interests  as  a  whole  and  thus  to  all  the 
people  of  the  state. 

The  western  meadowlark  has  an  esthetic  value  greater  than 
that  of  almost  any  other  bird  in  the  state.  This  value,  although 
not  capable  of  expression  in  dollars  and  cents,  is  nevertheless  real. 

Sufficient  protection  from  the  depredations  of  the  western 
meadowlark  is  afforded  the  farmers  of  the  state  through  a  law 
providing  for  the  "killing  of  a  meadowlark,  robin,  or  other  wild 
bird  by  the  owner  or  tenant  of  any  premises,  where  such  bird 
is  found  destroying  berries,  fruit  or  crops  growing  on  such 
premises. ' ' 

For  these  reasons,  the  western  meadowlark  (Sturnella  neg- 
lecta)  should  be  retained  on  the  list  of  protected  non-game  birds 
of  the  State  of  California  and  should  at  all  times  be  afforded  the 
protection  it  merits. 


1914]   Bryant:  Economic  Status  of  the  Western  Meadowlark     477 


SOME  INTERESTING  SIDE-LIGHTS  ON  THE 
INVESTIGATION 

In  an  investigation  of  this  kind  where  so  many  birds  of  the 
same  species  have  come  to  hand  it  would  be  unfortunate  if  some 
account  were  not  taken  of  certain  scientific  problems  such  as 
variation,  parasitism,  etc. 

PARASITISM 

Owing  to  the  fact  that  birds  were  received  in  the  laboratory 
preserved  in  formalin  it  has  been  impossible  to  examine  them 
for  blood  parasites.  The  only  parasites  discovered  have  been 
nematodes  (Spiroptera  sp.  ?),  which  have  been  found  in  the  body 
cavity  and  more  often  in  the  intestine.  Fewer  than  one  one- 
hundredth  of  one  per  cent  of  the  birds  examined,  however,  were 
found  to  be  infected.  Hence  western  meadowlarks  do  not  appear 
to  be  parasitized  to  any  great  extent  by  round  worms.  Western 
robins  have  shown  a  larger  percentage  of  infection. 

T.  G.  Pearson  (1909)  says  of  the  eastern  meadowlark:  "A 
parasitic  worm  often  found  in  the  small  of  the  back  deters  many 
experienced  persons,  however,  from  pursuing  the  bird  persist- 
ently." Whether  the  western  meadowlark  is  parasitized  to  a 
greater  or  less  extent  is  not  known,  owing  to  the  lack  of  data  on 
the  eastern  meadowlark  in  this  regard. 

Tachinid  larvae  have  been  taken  from  the  stomachs  of  mead- 
owlarks. As  these  larvae  are  common  parasites  of  grasshoppers 
and  crickets,  their  presence  in  the  stomachs  would  seem  to  be 
easily  explained.  However,  in  two  cases  at  least  they  were  eaten 
separately,  for  no  grasshoppers  or  crickets  were  found  in  the 
stomachs  containing  the  larvae. 

MALFORMATION 

Three  or  four  birds  received  in  the  laboratory  had  lost  a 
foot  or  leg.  In  each  instance  the  end  of  the  broken  tarsus  had 
become  enlarged  and  hardened  by  use.  One  bird,  handicapped 
by  the  loss  of  both  tarsi,  was  dwarfed. 


478          University  of  California  Publications  in  Zoology    [VOL.  11 

ALBINISM 

No  cases  of  albinism  in  the  meadowlarks  received  in  the 
laboratory  have  been  noted.  Variations  in  the  amounts  of  black, 
yellow,  and  white  markings  have  been  noted,  however.  Speci- 
mens received  from  the  northern  coast  region  have  been  the 
darkest  in  color,  those  from  the  southeastern  arid  regions  the 
lightest  in  color. 

ABUNDANCE  OF  THE  DIFFERENT  SEXES  AND  OF  JUVENILES 

If  the  numbers  of  the  sexes  collected  is  any  criterion  of  their 
comparative  numbers,  it  must  be  conceded  that  males  are  more 
abundant.  Fifty-eight  per  cent  of  one  thousand  birds  received 
from  ten  different  localities  were  males.  Collectors  usually  ob- 
tained a  larger  percentage  of  males  the  first  few  months.  This 
can  be  explained  on  the  grounds  that  the  male  is  a  larger  and 
more  conspicuous  bird.  Then,  too,  it  is  the  bird  most  often  seen 
during  the  nesting  season. 

About  seventy  per  cent  of  the  birds  collected  during  the 
summer  months  were  juveniles.  This  must  correspond,  to  some 
extent  at  least,  with  the  comparative  numbers  in  the  field. 


INCUBATION  AND  MOULT 

If  judged  by  the  condition  of  the  feathers  on  the  breast, 
females  do  most  of  the  incubation.  Breeding  males  were  noted 
as  early  as  late  February  and  as  late  as  August. 

None  of  the  birds  has  been  largely  denuded  of  feathers  even 
during  the  moulting  period.  At  no  time  do  they  appear  to  be 
greatly  hindered  by  moult. 


EFFECT  OF  SYSTEMATIC  DESTRUCTION  ON  NUMBERS 

A  systematic  destruction  of  birds  having  been  carried  on  in 
connection  with  this  investigation,  there  appeared  an  opportunity 
to  obtain  evidence  as  to  the  result  which  might  be  expected  as 
regards  a  decrease  in  numbers.  Consequently  a  letter  was  written 
to  deputies  enquiring  whether  there  had  been  a  perceptible  de- 
crease in  the  numbers  of  birds  in  the  localities  where  collections 


1914]   Bryant:  Economic  Status  of  the  Western  Meadoivlark     479 

were  made.  In  every  instance  where  a  reply  was  received  the 
deputy  reported  no  noticeable  decrease  in  numbers.  As  over 
two  hundred  western  meadowlarks  were  collected  during  the 
year  in  several  localities  it  would  seem  that  a  decrease  might  be 
in  evidence.  Of  course,  where  meadowlarks  are  abundant  such 
decrease  would  be  hard  to  detect.  However,  in  southern  Cali- 
fornia, where  meadowlarks  are  not  so  abundant,  it  seems  strange 
that  a  decrease  was  not  apparent. 

This  evidence  lends  support  to  th.e  view  that  the  meadowlark 
is  a  hardy,  prolific  bird  and  is  capable  of  withstanding  depletion 
far  better  than  certain  other  birds.  What  the  effect  of  placing 
the  bird  on  the  game  list  would  be  is  not  difficult  to  conjecture. 
The  continued,  systematic  destruction  which  such  a  move  would 
make  possible  would  certainly  have  a  greater  effect  on  numbers 
than  this  comparatively  slight  destruction  limited  to  not  more 
than  two  years. 

DEATH  RATE 

That  there  is  a  comparatively  small  percentage  of  young 
birds  which  grow  to  maturity  is  supported  by  the  fact  that 
during  a  two  weeks '  stay  in  a  region  where  western  meadowlarks 
were  nesting  abundantly,  three  dead  nestlings  were  found  at  dif- 
ferent times  and  in  addition  a  nest  of  four  destroyed  by  a  hawk 
or  weasel.  The  rearing  of  two  broods,  averaging  three  each,  also 
supports  this  view.  The  death  rate,  if  computed  from  the  average 
number  of  young  hatched,  would  be  seventy-five  per  cent,  for 
only  two  (twenty-five  per  cent)  out  of  every  eight  survive  if  the 
population  remains  the  same.  The  minimum  number  of  meadow- 
larks  can  be  expected  just  before  the  breeding  season.  As  the 
minimum  number  remains  fairly  constant  from  year  to  year,  it 
can  be  seen  that  from  every  pair  of  breeding  meadowlarks,  if 
they  lived  but  one  year,  it  would  be  possible  for  an  average 
maximum  of  only  two  to  reach  maturity.  Since  the  adults  must 
live  a  number  of  years  the  death  rate  must  be  greater  than 
seventy-five  per  cent. 

The  two  greatest  factors  in  the  death  rate  are  available  food 
supply  and  natural  enemies. 


480          University  of  California  Publications  in  Zoology    [VOL-  n 

Do  PROTECTIVE  ADAPTATIONS  OF  INSECTS  PROTECT  THEM  FROM 
THE  ATTACKS  OF  BIRDS? 

In  the  attempt  to  interpret  the  law  of  natural  selection 
emphasis  has  been  laid  on  the  "importance  of  any  structure  or 
character  which  enables  its  possessor  to  escape  destruction."  As 
soon  as  we  find  that  one  animal  preys  on  another,  we  immedi- 
ately seek  for  some  character  to  which  we  can  ascribe  the  sur- 
vival of  the  hunted  form.  This  has  led  to  an  overemphasis  on  the 
theory  of  protective  adaptations. 

Doubtless  the  interpretation  we  have  put  upon  certain  color 
characters  and  other  characters  called  protective  have  been  of 
value,  just  as  our  imperfect  systems  of  classification  have  been 
of  value.  Yet,  just  as  we  are  constantly  changing  and  modi- 
fying original  classifications,  so  we  may  expect  to  modify  our 
views  concerning  protective  adaptations. 

In  order  to  point  out  the  view  usually  held,  attention  is  called 
to  the  following  quotation  from  Kellogg  (1908)  :  "It  has  been 
conclusively  shown  by  observation  and  experiment,  by  several 
naturalists,  that  many  insects  are  distasteful  to  birds,  lizards 
and  other  enemies  of  the  insect  class.  The  blood,  lymph  or  some 
specially  secreted  body  fluid  of  these  insects  contains  an  acrid  or 
ill-tasting  substance,  so  that  birds  will  not,  if  they  can  recog- 
nize the  kind  of  insect,  make  any  attempt  to  catch  or  eat  them. ' ' 

Kellogg  also  goes  on  to  suggest  the  theory  that  "success"  is 
dependent  on  protective  adaptations.  Certain  animals  are 
widespread  and  found  in  great  numbers  and  certain  others  in 
small  numbers.  Those  existing  in  great  numbers  are  said  to  do 
so  because  they  are  protected  from  their  enemies.  Enemies  are 
only  one  factor  in  the  complex  that  governs  the  abundance  or 
scarcity  of  a  species,  so  that  such  a  theory  hardly  seems  justified. 

The  examination  of  so  large  a  number  of  stomachs  of  one 
species  of  bird  has  furnished  some  interesting  evidence  regarding 
the  extent  to  which  certain  insects  are  protected  from  their  ene- 
mies. The  evidence  shows  that  many  of  the  so-called  protective 
adaptations  of  insects  do  not  protect  them  from  the  attacks  of 
enemies  to  the  extent  to  which  we  have  been  led  to  believe. 

A  recent  paper  by  W.  L.  McAtee  (1912d)  of  the  U.  S.  Biolog- 
ical Survey,  entitled  "The  experimental  method  of  testing  the 


1914]   Bryant:  Economic  Status  of  the  Western  Meadowlark     481 

efficiency  of  warning  and  cryptic  coloration  in  protecting  animals 
from  their  enemies, ' '  clearly  points  out  the  fact  that  the  tests  of 
protective  adaptations  against  natural  enemies  have  been  incon- 
sistent, misinterpreted,  and  are  untrustworthy  guides  to  behavior 
under  natural  conditions. 

Investigators  using  the  experimental  method  have  too  often 
failed  to  take  into  account  other  factors  instrumental  in  modi- 
fying the  behavior  of  animals  toward  their  prey.  "The  rejection 
of  various  items  of  food  by  captive  animals  does  not  prove  that 
these  items  are  rejected  by  the  same  species  under  natural  con- 
ditions." It  does  give  some  idea  of  the  food  habits,  but  does 
not  furnish  dependable  evidence  as  to  the  food  of  birds  in  the 
wild. 

More  reliance  can  be  placed  on  the  evidence  furnished  by 
stomach  examination,  for  "there  is  no  possibility  of  going  back 
of  such  evidence  on  the  choice  of  food."  Its  one  drawback  is 
that  it  does  not  furnish  us  with  data  as  to  what  was  not  chosen. 

Hence  the  following  evidence  must  be  considered  as  valuable 
in  throwing  light  upon  this  much  discussed  problem.  The  dis- 
cussion will  be  largely  directed  to  such  protective  adaptations  as 
stings,  noxious  secretions,  hairs,  etc.,  as  the  evidence  at  hand 
bears  more  directly  on  this  phase  of  the  subject. 

More  has  been  written  on  the  palatability  of  butterflies  than 
on  any  other  insect.  To  back  up  the  theory  of  mimicry  it  was 
necessary  that  birds  be  made  an  important  enemy  of  butterflies. 
That  birds  are  an  important  enemy  of  butterflies  still  remains 
to  be  proved.  The  fact  that  the  records  of  the  United  States 
Biological  Survey  show  that  in  the  examination  of  40,000  stom- 
achs of  birds  but  four  cases  have  been  found  where  the  birds 
concerned  had  eaten  butterfles  would  support  Mr.  McAtee's 
contention  that  "butterflies  are  in  very  little  demand  with  birds 
in  the  United  States."  On  the  other  hand,  the  fact  that  eleven 
butterflies  have  been  taken  from  the  stomachs  of  western  meadow- 
larks  and  that  five  different  species  of  birds  were  found  feeding 
on  butterflies,  to  a  greater  or  less  extent,  during  an  outbreak  of 
these  insects  in  northern  California  during  the  spring  and  sum- 
mer of  1911  (Bryant,  1911),  shows  that  butterflies  are  taken  to 
some  extent  as  food.  The  observational  evidence  of  Mr.  Tyler 


482          University  of  California  Publications  in  Zoology    [VOL.  11 

already  quoted  (p.  430)  also  supports  the  latter  statement.  The 
ten  pages  of  evidence  given  by  Poulton  (1908)  also  supports  the 
view  that  birds,  even  though  they  do  not  feed  to  any  considerable 
extent  on  these  insects,  do,  occasionally  at  least,  feed  upon  them. 
Mason  and  Lefroy  (1912),  after  a  study  of  the  food  of  the  birds 
of  India,  state:  "Butterflies  do  not  form  any  appreciable  pro- 
portion of  the  food  of  any  one  species  of  bird,  though  a  good 
many  birds  take  these  insects  at  times. ' ' 

Although  identification  of  the  butterflies  found  in  the  stom- 
achs has  often  been  impossible,  yet  it  is  certain  that  they  often 
belonged  to  separate  families.  The  difference  between  the  pierid 
butterfly,  Eurymus  eurytheme,  and  the  nymphalid  butterfly, 
Eugonia  calif ornica,  is  great  enough  to  show  that  there  is  little 
choice  shown.  This  evidence  is  supported  by  the  work  of  Manders 
(1911).  He  concludes:  "There  is  no  bird  in  Ceylon  known  to 
eat  butterflies  that  distinctly  discriminates  as  an  adult  between 
one  species  of  butterfly  and  another."  Manders  also  goes  so  far 
as  to  say :  '  *  The  fact  that  there  is  no  discrimination  shown  by 
adults  leads  one  to  conclude  either  that  few  or  no  tasting  experi- 
ments were  undertaken  in  youth,  or,  what  is  more  probable,  that 
their  taste  with  regard  to  them  is  indifferent."  One  important 
criticism  that  can  be  made  of  Mander  's  work  is  that  he  depended 
entirely  on  observation.  The  criticism  bears  less  weight,  how- 
ever, since  the  insects  under  observation  were  of  sufficient  size 
and  conspicuousness  to  make  the  observational  method  more 
dependable. 

Any  one  who  has  watched  a  bird  catch  a  butterfly  must 
necessarily  be  impressed  with  the  skill  needed.  Certain  agility 
on  the  part  of  the  butterfly  must  aid  greatly  in  protecting  it 
from  attack.  Further  study  may  demand  a  change  of  emphasis 
from  protective  coloration  to  the  protection  afforded  by  the  but- 
terfly's ability  to  elude  its  pursuer. 

There  appears  to  be  evidence  that  birds  seldom  attack  butter- 
flies, thus  lending  support  to  the  theory  that  they  are  protected. 
On  the  other  hand,  it  can  be  seen  that  there  is  evidence  that 
butterflies  are  taken  regularly  as  food  by  some  birds.  There 
are  doubtless  many  factors  which  enter  into  the  problem  which 
have  not  as  vet  been  considered.  Hence  until  further  evidence 


1914]   Bryant:  Economic  Status  of  the  Western  Meadowlark     483 

is  forthcoming  the  best  conclusion  would  seem  to  be  that  birds 
occasionally  eat  butterflies,  and  that  when  they  become  extremely 
available  the  number  taken  is  increased. 

Spiny  and  hairy  caterpillars  have  often  been  pointed  out  as 
specially  well  protected  insects.  At  least  fifty  of  the  stomachs 
examined  have  contained  these  caterpillars.  In  one  instance  the 
larva  of  the  mourning-cloak  butterfly  (Euvanessa  antiopa)  has 
been  taken  from  a  stomach.  The  size  as  well  as  the  spiny  char- 
acter of  this  caterpillar  would  seem  to  preclude  attack.  Numbers 
of  small  hairy  caterpillars  have  been  taken  from  the  stomachs. 
Judd  (1899)  states  that  "The  hairiness  of  caterpillars  seems  to 
secure  them  from  the  attack  of  birds  more  effectually  than  do 
any  of  the  protective  devices  so  far  considered."  Comparing 
the  relative  abundance  of  these  caterpillars  available  for  the 
western  meadowlark  with  those  not  so  protected,  it  seems  safe  to 
say  that  certain  of  the  smaller  hairy  caterpillars  are  not  often 
passed  by  because  of  their  hairiness. 

Stink-bugs  (Pentatomidae),  in  spite  of  their  noxious  secretion 
and  disagreeable  odor,  form  a  constant  article  of  diet  for  the 
western  meadowlark.  In  the  examination  of  a  collection  of  birds 
from  Newman,  Stanislaus  County,  it  was  found  that  stink-bugs 
(Euschistus,  Podisus,  Alydus,  Coryzus)  had  been  taken  every 
month  from  March  to  October,  inclusive,  and  formed  five  and 
three-tenths  per  cent  of  the  food  for  the  year.  Pinicate  beetles 
(Eleodes  sp.),  having  a  noxious  secretion,  are  commonly  taken 
as  food  by  meadowlarks. 

Many  stinging  insects  also  form  a  constant  article  of  diet. 
Chief  among  these  are  ants,  bees  and  wasps,  and  cow-killers 
(Mutillidae).  Kissing-bugs  (Reduviidae)  have  been  found  in  a 
few  instances.  The  stomachs  of  two  out  of  four  birds  eating 
reduviids  were  empty,  indicating  that  the  poison  might  have 
caused  some  discomfort.  Bees  and  wasps  are  so  often  taken  that 
it  can  hardly  be  said  that  their  stinging  propensities  preclude 
attack.  Probably  their  agility  is  much  more  important  in  pro- 
tecting them  from  the  attack  of  birds.  Over  two  hundred  ants 
have  been  taken  from  a  single  stomach.  If  the  stings  or  the 
poison  had  any  effect,  it  does  not  seem  reasonable  that  a  bird 
would  feed  exclusively  on  ants  even  when  hard  pressed  for  food. 


484          University  of  California  Publications  in  Zoology    1T°L-  H 

The  finding  of  scorpions  in  the  stomachs  of  two  meadowlarks 
from  San  Diego  was  a  surprise  for  two  reasons.  Their  size  and 
sting  would  apparently  protect  them.  In  addition,  their  noc- 
turnal habits  and  their  habitat  would  seem  to  offer  protection. 
That  more  were  not  taken  is  probably  due  to  the  fact  that  they 
are  seldom  available  rather  than  that  they  are  unpalatable. 

The  investigation  has,  therefore,  shown  that  insects  suppos- 
edly protected  by  noxious  secretions,  malodor,  stings,  etc.,  are 
taken  as  food  by  western  meadowlarks.  According  to  the  old 
idea,  the  survival  of  these  insects  can  be  traced  directly  to  pro- 
tective adaptations.  Earty  authors  even  suggested  almost  com- 
plete immunity  from  attack.  In  recent  years  this  view  has  become 
modified.  Judd  (1899)  called  attention  to  the  fact  that  "Biol- 
ogists have  not  yet  entirely  elucidated  all  the  details  of  the  nature 
of  adaptations  of  insects  which  are  potently  protected."  The 
same  thing  can  be  said  at  the  present  time. 

If  we  hold  to  the  theory  of  natural  selection,  it  is  important 
that  a  certain  toll  be  taken  in  order  to  perfect  adaptation.  If 
an  insect  had  no  enemies  it  would  have  little  need  of  protective 
adaptations.  Of  course  it  may  be  argued  that  after  the  adapt- 
ation becomes  perfected  the  enemies  of  insects  learn  to  let  them 
alone.  Unless  the  variations  were  of  the  orthogenetic  type,  how- 
ever, we  could  hardly  expect  such  highly  differentiated  protective 
adaptations  to  exist  as  do  exist.  Kellogg 's  view  that  "success" 
is  dependent  on  protective  adaptations  rests  on  this  assumption. 
It  overemphasizes  the  part  played  by  protective  adaptations.  In 
the  working  of  the  principle  of  natural  selection  other  principles 
and  tendencies  are  working  against  the  factor  of  protective 
adaptations,  and  it  cannot  be  said  that  protective  adaptations 
gain  the  ascendency  over  all  other  tendencies. 

Unusual  destruction  of  so-called  protected  insects  and  other 
arthropods  can  sometimes  be  attributed  to  young  birds.  Finn 
(1898)  found  out  by  experiment  "that  each  bird  has  to  separately 
acquire  its  experience,  and  well  remembers  what  it  has  learned. 
Lloyd  Morgan  (1896)  has  also  shown  that  birds  have  no  instinc- 
tive knowledge  of  the  different  kinds  of  food,  but  that  they 
examine  and  test  everything.  He  also  points  out  the  fact  that 
they  have  excellent  memories  and  are  able  to  remember  sum"- 


1914]   Bryant:  Economic  Status  of  the  Western  Meadowlark     485 

ciently  so  well  any  unpleasant  sensation  that  they  usually  avoid 
a  recurrence. 

Under  such  circumstances,  we  should  naturally  expect  that 
most  of  the  birds  taking  protected  insects,  etc.,  would  be  young 
and  inexperienced  birds.  In  the  present  investigation  Juvenal 
birds  have  been  differentiated  from  adults  as  far  as  possible. 
Consequently  evidence  on  this  point  is  available.  Contrary  to 
expectation,  most  of  the  birds  taking  protected  insects  and  other 
arthropods  as  food  have  been  adult  birds.  Their  previous  ex- 
perience with  this  kind  of  food  is  unknown.  The  simplest  expla- 
nation is  to  say  that  birds  in  searching  for  food  take  that  at  hand 
and  that  most  easily  obtained. 

The  only  evidence  afforded  by  this  investigation  that  birds 
learn  to  let  certain  insects  alone  is  the  total  lack  of  cocci  nellid 
beetles  in  the  food  of  the  western  meadowlark.  In  California 
at  certain  times  of  year  coccinellid  beetles  are  extremely  common 
and  certainly  would  form  an  available  food  for  the  meadowlark 
if  they  were  not  protected  from  attack.  The  chrysomelid  beetle, 
Diabrotica  soror,  a  beetle  sometimes  confused  with  coccinellids, 
is  occasionally  taken  as  food.  One  stomach  was  found  completely 
filled  with  these  beetles.  Hence  it  would  seem  that  birds  can 
distinguish  between  coccinellids  which  appear  to  be  noxious  and 
certain  chrysomellids  which  appear  to  be  edible.  They  also  dis- 
tinguish between  pentatomids  with  a  noxious  secretion  and  cocci- 
nellids with  a  similar  secretion. 

Movement  is  a  very  important  factor  connected  with  the 
problem.  Allen  (1912)  and  Roosevelt  (1911)  have  emphasized 
this  point  of  view,  that  any  coloration  is  protective  only  so  long 
as  the  animal  is  motionless.  Allen  goes  so  far  as  to  say  that 
"coloration  is  a  minor  asset  in  an  animal's  protection  in  com- 
parison with  its  other  qualities — alertness,  truculence  and  other 
traits  that  make  for  its  protection."  This  point  of  view  has 
received  far  less  emphasis  than  its  importance  justifies. 

The  highest  expression  of  vision  is  to  be  found  in  birds.  The 
color  sense,  especially,  is  very  acute,  as  shown  by  the  preponder- 
ence  of  cones  in  the  retinal  elements.  The  range  and  rapidity 
of  accommodation  in  birds  far  exceeds  that  of  man  or  other 
animals,  and  the  accommodative  and  refractive  apparatus  is  much 


486          University  of  California  Publications  in  Zoology    [VOL.  11 

more  complex  than  in  the  other  subkingdoms  (Wood,  1907). 
Birds  must,  therefore,  be  able  to  distinguish  readily  the  differ- 
ently colored  insects  as  well  as  to  note  quickly  differences  in  size 
and  any  motion  on  their  part.  Hence  the  food  taken  must  be 
largely  the  result  of  the  reflex  acts  set  up  by  the  sight  of  food 
and  much  less  as  the  result  of  studying  each  kind  of  food  pre- 
sented to  determine  its  palatability.  This  brings  us,  of  course, 
to  "hunger"  as  one  of  the  prime  determining  agents  as  to  the 
kind  as  well  as  the  amount  of  food  taken. 

Then,  too,  many  insects  and  animals  are  protectively  colored 
while  at  rest,  but  let  them  move  and  they  immediately  become 
conspicuous.  Experiment  has  shown  that  most  of  the  lower  verte- 
brates depend  largely  on  the  movement  of  their  prey  to  apprize 
them  of  its  presence.  A  fly  placed  in  a  cage  with  horned  lizards 
is  unnoticed  until  it  moves.  Many  a  protectively  colored  insect 
must  escape  detection  because  it  remains  at  rest.  Let  the  same 
insect  move  and  it  is  instantly  detected. 

Let  us  take  a  few  examples  in  the  food  of  the  western  meadow- 
lark.  Many  of  the  snout-beetles  (Otiorhynchidae)  are  inconspic- 
uously colored  and  often  so  covered  with  dust  as  to  be  the  exact 
color  of  the  ground.  Close  search  often  fails  to  disclose  a  grass- 
hopper on  a  grass-stem  or  weed,  so  well  does  it  blend  with  its 
surroundings.  The  same  can  be  said  of  stink-bugs.  Yet  all  of 
these  insects  are  taken  in  large  numbers  by  meadowlarks.  The 
explanation  probably  lies  in  this  factor  of  movement.  These 
insects,  although  well  concealed  while  at  rest,  are  not  concealed 
when  moving,  but  are,  on  the  contrary,  conspicuous. 

An  insect  outside  of  its  own  environment  is  also  easily  de- 
tected by  its  enemies.  A  stink-bug,  although  inconspicuous  on 
a  green  plant  stem,  becomes  conspicuous  on  the  bare  ground. 
This  again  furnishes  a  possible  reason  for  the  large  numbers  of 
these  insects  found  in  the  stomachs  of  western  meadowlarks. 

Movement  and  particular  environment  modify  the  value  of 
protective  or  concealing  coloration.  There  may  also  be  still  other 
factors  which  modify  its  value.  Size  and  bright  coloration  must 
add  to  an  insect 's  conspicuousness  in  both  movement  and  change 
of  environment.  All  evidence  from  this  investigation  points  to 
the  fact  that  although  certain  insects  may  be  protectively  colored, 


1914]   Bryant:  Economic  Status  of  the  Western  Meadowlark     487 

yet  they  are  not  immune  from  the  attack  of  birds  because  of 
such  coloration.  The  fact  that  protective  coloration  is  of  maxi- 
mum utility  to  the  insect  only  when  it  is  at  rest,  and  of  minimum 
utility  when  it  is  moving  or  when  it  is  out  of  its  natural  habitat, 
in  a  large  measure  explains  the  occurrence  of  these  insects  in 
the  food  of  birds. 

Cow-killers  (Mutillidae)  have  been  found  in  several  instances. 
These  insects  are  usually  considered  as  being  warningly  colored. 
Were  they  as  abundant  as  other  insects  the  numbers  taken  by 
western  meadowlarks  would  be  insignificant.  In  that  they  are 
not  numerous,  the  fact  that  five  stomachs  contained  them  indi- 
cates that  their  warning  coloration  did  not  protect  them  wholly 
from  attack. 

It  may  even  be  said  that  certain  unpalatable  insects  are  taken 
as  food  simply  because  they  are  made  conspicuous  by  movement, 
thus  setting  up  a  chain  of  reflexes  in  the  bird  which  result  in 
their  being  eaten.  The  reflexes  set  up  by  the  stimulus  of  the 
sight  of  food  play  an  important  part  in  determining  the  kind 
and  amount  of  food  taken.  A  bird  feeding  on  grasshoppers 
would  doubtless  be  more  greatly  influenced  by  the  sight  of  another 
grasshopper  than  by  that  of  a  small  beetle  or  even  a  cricket  the 
same  size  as  the  grasshopper.  The  psychological  process  involved 
in  the  feeding  habit  has  been  little  studied.  Its  importance  as 
a  factor  suggests  this  as  a  fruitful  source  from  which  might  come 
illuminating  evidence  on  the  problem. 


AVAILABILITY  AS  A  FACTOR  IN  THE  KIND  AND  QUANTITY  OF  FOOD 
Western  meadowlarks  collected  in  grain  fields  appear  to  take 
as  food  practically  every  kind  of  insect  and  other  arthropod  to  be 
found  in  grain  fields.  Small  size  does  not  govern  the  kind  of 
food,  for  one  bird  was  found  to  have  eaten  aphids.  Nor,  on  the 
other  hand,  does  large  size  preclude  attack,  for  pinicate  beetles 
(Eleodes  sp.)  are  eaten.  Insects  with  stings,  such  as  ants,  bees, 
and  wasps,  insects  with  noxious  secretions,  such  as  stink-bugs, 
and  even  hairy  caterpillars  are  regularly  taken  as  food.  The 
respective  quantity  of  each  kind  taken  appears  to  parallel  their 
abundance  and  accessibility.  The  term  availability  denotes  the 


488          University  of  California  Publications  in  Zoology    [VOL.  11 

totality  of  factors  governing  the  food-taking  of  birds.  In  other 
words,  in  spite  of  the  fact  that  birds  select  food  to  a  greater 
extent  than  many  other  animals,  they  do  not  select  kinds  or 
quantities  of  food  elements  to  the  degree  to  which  we  have  been 
led  to  believe  from  experiments  on  captive  birds,  but  are  gov- 
erned more  largely  by  the  abundance,  the  ease  of  capture,  the 
conspicuousness,  etc.,  of  the  insect,  or,  in  other  words,  its  avail- 
ability. 

The  attempt  has  been  made  heretofore  to  interpret  the  food 
habits  of  birds  from  the  standpoint  of  the  insect,  an  endeavor 
being  made  to  show  why  certain  insects  were  not  taken  as  food. 
Looking  at  the  same  problem  from  the  standpoint  of  the  bird, 
we  can  say  that  with  few  exceptions  the  term  availability,  when 
defined  as  above,  best  expresses  the  interrelations  associated 
with  the  food  habits  of  birds.  Under  this  term  can  be  grouped 
both  the  objective  factors  concerned,  such  as  abundance,  ease  of 
capture,  conspicuousness,  etc.,  and  the  psychological  factors 
involved  in  the  taking  of  food. 

SOLVED  AND  UNSOLVED  PROBLEMS  IN  ECONOMIC 
ORNITHOLOGY 

In  spite  of  the  advance  made  in  the  study  of  the  food  of  birds 
and  their  relation  to  agriculture,  there  are  still  many  problems 
connected  with  economic  ornithology  that  are  still  unsolved. 

Great  improvement  in  the  technique  of  determining  the  food 
of  birds  has  been  achieved.  The  modern  method  of  stomach 
examination  combined  with  field  observation  is  so  far  ahead  of 
the  former  method  of  simple  observation  or  inferential  reasoning 
that  one  is  led  to  believe  that  a  solution  of  many  of  the  problems 
presented  is  near  at  hand.  The  criteria  used  in  determining  the 
status  of  a  bird  have  also  been  improved  to  such  an  extent  that 
were  it  possible  to  obtain  the  prescribed  kind  of  evidence  our 
estimates  of  the  value  of  a  bird  must  needs  be  near  the  truth. 

Our  goal,  "a  balance  of  all  the  benefits  conferred  against  all 
the  injuries  inflicted,"  is  a  fine  ideal,  but  we  are  still  unable  to 
attain  it.  A  great  deal  is  still  made  of  the  fact  that  certain  birds 
eat  large  quantities  of  injurious  insects.  Yet  what  are  injurious 
insects?  Insects  injurious  in  one  place  may  be  comparatively 


1914]   Bryant:  Economic  Status  of  the  Western  Meadowlark     489 

unimportant  in  another  place.  Abundance  even  more  than  the 
kind  of  an  insect  appears  to  govern  its  injuriousness.  Hence 
the  direct  value  of  the  destruction  of  certain  insects  by  birds  is 
difficult  to  estimate. 

The  solution  of  the  problem  lies  in  a  determination  of  the 
comparative  abundance  of  insects.  The  censuses  of  birds  taken 
by  Forbes  (1901)  in  Illinois  have  been  of  incalculable  value  to 
ornithologists  and  everyone  regrets  the  lack  of  others  of  the  same 
type.  The  entomologist  has  left  this  quantitative  phase  almost 
wholly  untouched.  Consequently  the  economic  ornithologist,  in 
attempting  to  determine  the  good  accomplished  by  birds  in  de- 
stroying insects,  has  no  evidence  on  which  he  can  depend  as  to 
the  comparative  numbers  of  the  insects  upon  which  they  feed. 
The  difficulty  of  obtaining  any  exact  idea  of  the  numbers  of  a 
species  of  insect  in  the  field  is  very  great,  but  the  importance 
and  value  of  the  information  will  certainly  repay  effort  made 
in  this  direction.  Economic  ornithologists  must  necessarily  await 
data  of  this  kind  before  they  will  be  able  to  point  out  conclusively 
the  importance  to  be  attached  to  the  destruction  of  insects  by 
birds. 

In  years  past  we  have  been  wont  to  judge  the  value  of  a  bird 
entirely  on  its  food.  The  present  tendency  to  regard  the  food 
habit  as  only  one  of  the  things  to  be  considered  in  judging  its 
value  is  an  advance  worthy  of  note.  The  esthetic  value  is  fast 
coming  to  be  appreciated  by  every  one.  It  must  certainly  be 
given  an  important  place  in  any  adequate  estimate  of  a  bird's 
value. 

In  a  broad  sense  food  preference  furnishes  evidence  as  to  the 
value  of  a  bird.  For  instance,  a  bird  that  eats  insects  can  be 
considered  of  more  value  to  the  agriculturist  than  one  that  sub- 
sists entirely  on  vegetable  food.  To  particularize  and  say  that 
a  bird  prefers  a  certain  kind  of  insect  is  to  tread  on  dangerous 
ground,  for  birds  appear  to  be  governed  more  largely  by  the 
abundance  of  an  insect  than  by  its  taste.  The  opportunity 
afforded  a  bird  for  obtaining  an  insect  appears  to  be  a  stronger 
factor  than  memory  of  a  bad  taste.  Consequently  the  old  idea 
of  food  preference  must  be  modified  to  meet  the  modern  idea  of 
availability  as  a  factor  in  the  kind  and  quantity  of  food. 


490          University  of  California  Publications  in  Zoology    [VOL.  ll 

The  exact  relation  which  birds  bear  to  the  control  of  insect 
outbreaks  remains  to  be  worked  out.  There  are  a  number  of 
factors  which  tend  to  bring  back  normal  conditions.  Whether 
birds,  parasitic  insects,  weather  conditions,  or  other  factors  are 
of  prime  importance  is  a  problem  yet  to  be  solved.  The  begin- 
ning has  been  made  and  significant  evidence  adduced.  A  compre- 
hensive study  of  this  problem  would  do  much  towards  solving  it. 

The  amount  of  benefit  to  be  derived  through  the  destruction 
of  weed  seeds  is  still  a  debated  question.  When  such  efficient 
measures  as  the  weed  cutter  and  the  mowing  machine  can  be 
brought  into  use,  the  amount  of  good  accomplished  in  a  bird's 
destruction  of  weed  seeds  is  minimized.  A  weed-seed  eater  must 
be  considered  a  less  valuable  bird  than  an  insect  eater  where 
weeds  are  more  easily  controlled  than  insects.  Better  insect 
control  measures  will  likewise  make  the  value  of  insectivorous 
birds  less  apparent.  The  inherent  value,  nevertheless,  remains 
unchanged.  Common  sense  would  seem  to  dictate  the  making 
use  of  natural  control  measures  to  the  greatest  extent  possible. 
Herein  lies  the  justification  for  emphasis  on  the  value  of  birds 
as  insect  and  weed-seed  destroyers. 

SUMMARY 

Owing  to  the  constant  complaint  of  ranchers  as  to  the  depre- 
dations of  birds  and  attempted  legislation  to  take  protection 
away  from  certain  non-game  birds,  the  California  State  Fish  and 
Game  Commission,  in  co-operation  with  the  University  of  Cali- 
fornia, has  undertaken  a  thorough,  scientific  investigation  into 
the  relations  of  birds  to  agricultural  interests.  The  western 
meadowlark  (Sturnella  neglect  a),  owing  to  its  depredations  in 
sprouting  grain  fields,  has  been  the  first  bird  to  be  studied. 

A  study  of  the  history  of  methods  used  in  economic  ornith- 
ology has  shown  that  the  time  has  come  when  circumstantial  or 
partial  evidence  as  to  the  food  of  a  bird  is  insufficient  evidence 
on  which  to  determine  its  economic  status.  Nothing  short  of  a 
knowledge  of  a  bird's  food  for  the  whole  year,  a  knowledge  of 
its  depredations,  and  its  whole  life  history,  allowing  a  balance 
of  all  of  the  benefits  conferred  against  all  of  the  injuries  done, 
is  now  demanded. 


1914]   Bryant:  Economic  Status  of  the  Western  Meadowlark     491 

The  western  meadowlark  is  distributed  throughout  the  state, 
most  abundantly  in  the  great  interior  valleys,  the  grain-growing 
districts.  It  appears  to  be  increasing  in  numbers  in  localities 
where  cultivated  crops  are  furnishing  it  better  food  and  cover. 

The  investigation  has  included  field  investigation  of  birds  at 
the  field  of  action,  experimentation  on  the  amounts  of  food  con- 
sumed and  the  times  of  digestion,  and  stomach  examinations  of 
the  kinds  and  quantities  of  food  actually  consumed. 

Over  twenty  collections  from  different  parts  of  the  state, 
made  up  of  birds  collected  each  month  of  the  year  in  the  same 
general  locality,  have  been  available  for  stomach  examination. 
Nearly  twro  thousand  stomachs  of  western  meadowlarks  have  been 
examined  and  the  kind  and  quantity  of  the  different  elements  of 
food  tabulated. 

Field  investigation  has  shown  that  the  western  meadowlark 
destroys  sprouting  grain  by  boring  down  beside  the  sprout  and 
eating  the  kernel.  The  amount  of  damage  depends  upon  the 
abundance  of  the  birds,  the  depth  to  which  the  grain  is  planted, 
the  size  of  the  field,  the  condition  of  the  soil,  the  proximity  to 
pasture  or  uncultivated  land,  and  the  time  of  year.  The  amount 
of  damage  possible  is  minimized  by  the  short  time  (two  weeks) 
during  which  damage  to  the  young  plant  can  result.  No  other 
crops  are  seriously  damaged  by  western  meadowlarks.  Young 
meadowlarks  are  fed  exclusively  on  insect  food,  principally  cut- 
worms and  grasshoppers. 

Experimentation  on  captive  birds  has  shown  that  nestling 
birds  consume  very  nearly  their  own  weight  of  food  every  day. 
The  time  during  which  insects  remain  in  the  stomach  is  from 
three  to  four  hours.  The  time  during  which  grain  remains  in 
the  stomach  is  from  four  to  six  hours.  Hence  the  time  of  diges- 
tion of  grain  is  longer  than  that  of  insects.  The  amount  of 
insect  food  found  in  the  stomach  of  a  western  meadowlark  repre- 
sents, therefore,  but  one-third  of  the  daily  requirement. 

Stomach  examination  has  shown  that  sixty-three  and  three- 
tenths  per  cent  of  the  total  volume  of  food  of  the  western  meadow- 
lark  for  the  year  is  made  up  of  animal  matter  and  thirty-six  and 
seven-tenths  per  cent  of  vegetable  matter.  The  animal  matter 
is  made  up  mostly  of  ground  beetles,  grasshoppers,  crickets,  cut- 


492          University  of  California  Publications  in  Zoology    [VOL. 11 

worms,  caterpillars,  wireworms,  stink-bugs,  and  ants,  insects  most 
of  which  are  injurious  to  crops.  The  vegetable  matter  is  made 
up  of  grain  and  seeds.  Grain  as  food  reaches  a  maximum  in 
November,  December,  and  January,  insects  in  the  spring  and 
summer  months,  and  weed  seeds  in  September  and  October.  An 
average  stomach  of  a  western  meadowlark  contains  two  and 
three-fourths  cubic  centimeters  of  food.  The  kind  and  quantity 
of  food  varies  with  the  time  of  year  and  locality. 

A  study  of  the  relation  of  birds  to  insects  has  shown  that 
birds  are  important  regulators  of  the  numbers  of  insects  for  at 
least  two  reasons :  ( 1 )  the  maximum  consumption  of  insects  comes 
during  the  nesting  season  of  birds,  a  time  when  the  numbers  of 
insects  are  at  a  maximum;  (2)  birds  change  their  food  habits 
and  feed  on  the  insect  most  available,  thereby  becoming  important 
balancers  during  insect  outbreaks  when  insects  appear  in  ab- 
normal numbers. 

The  verdict  of  ranchers  throughout  the  state  obtained  by  a 
circular  letter  has  shown  that  there  is  a  wide  difference  of  opinion 
as  to  the  extent  of  damage  caused  by  the  western  meadowlark. 
A  majority  maintain  that  this  bird  does  not  damage  crops  and 
is,  therefore,  not  a  nuisance. 

A  comparison  of  the  injuries  caused  by  the  western  meadow- 
lark  with  the  benefits  it  confers  shows  that  it  does  more  good 
than  harm,  and  so  merits  protection  as  an  insectivorous  non-game 
bird.  Crops  may  be  largely  protected  by  drilling  grain  deeply 
or  by  furnishing  the  birds  sufficient  food  by  heavier  sowing. 

The  examination  of  so  large  a  number  of  birds  of  one  species 
has  furnished  some  interesting  side-lights  on  the  investigation. 
Evidence  on  such  biological  problems  as  parasitism,  malforma- 
tion, albinism,  natural  death  rate,  etc.,  has  been  made  available. 

Protective  adaptations  of  insects  do  not  render  them  immune 
from  the  attack  of  birds.  This  investigation  has  demonstrated 
the  following  facts  bearing  on  the  relation  of  birds  to  insects 
with  protective  adaptations: 

1.  Protective  adaptations  of  arthropods  such  as  stings,  nox- 
ious secretions,  hairs,  etc.,  have  been  overemphasized  as  factors 
protecting  the  owners  from  the  attack  of  birds. 

2.  The  "availability"  of  an  insect  or  arthropod,  when  this 
term  is  made  to  include  the  totality  of  factors  governing  the 


1914]   Bryant:  Economic  Status  of  the  Western  Meadowlark     493 

taking  of  an  insect  by  a  bird,  can  be  considered  the  most  potent 
factor  governing  the  food  habits  of  birds. 

3.  Hunger  or  inexperience  fails  to  account  for  the  destruction 
of    many  of    the  so-called  protected  insects,   because  in  many 
instances  they  form  staple  articles  of  diet. 

4.  The  fact  that  coccinellid  beetles  almost  wholly  escape  the 
attack  of  birds  is  the  one  thing  that  supports  the  theory  that 
certain  arthropods  are  practically  immune  from  the  attack  of 
birds  in  spite  of  the  fact  that  they  are  available  as  food.     The 
logical  explanation  of  the  immunity  lies  in  the  possession  of  a 
noxious  secretion  which  makes  them  unpalatable.    Why  this  secre- 
tion should  be  more  effective  than  that  of  the  Pentatomidae  or 
certain  tenebrionid  beetles  is  not  known. 

5.  Insects  protectively  colored  when  at  rest  are  easily  detected 
by  birds  or  other  enemies  as  soon  as  they  move.    An  insect  out- 
side of  its  natural  habitat  also  becomes  easy  to  detect.     These 
factors  of  movement  and  change  of  habitat  may  explain  the 
occurrence  of  certain  protectively  colored  insects  in  the  diet  of 
the  western  meadowlark. 

6.  The  reflexes  set  up  by  the  stimulus  of  the  sight  of  food, 
or,  in  other  words,  the  psychological  processes  involved  in  the 
taking  of  food,  doubtless  play  an  important  part  in  the  kind  and 
quantity  of  food  taken. 

A  number  of  factors  govern  the  kind  and  amount  of  food 
taken  by  birds.  The  totality  of  such  factors  (e.g.,  abundance, 
palatability,  ease  of  capture,  conspicuousness,  etc.)  can  be  ex- 
pressed by  the  term  availability.  Availability,  thus  used,  best 
accounts  for  the  varying  food  habits  of  birds. 

Although  great  progress  has  been  made  in  economic  ornith- 
ology, there  are  still  a  number  of  problems  that  remain  unsolved, 
chief  of  which  is  the  value  to  be  placed  on  the  destruction  of 
insects  and  weed  seeds  effected  by  birds.  Since  a  solution  of 
this  problem  depends  upon  quantitative  studies  of  the  compar- 
ative abundance  of  different  species  of  insects  and  weed  seeds, 
economic  ornithology  must  await  such  studies  from  the  entomol- 
ogist and  botanist. 

Transmitted  May  2,  1913. 


494          University  of  California  Publications  in  Zoology    [VOL.  11 


BIBLIOGRAPHY 

N.  B. — For  papers  on  economic  ornithology  previous  to  1901  see  Weed 
(1902).  For  publications  of  the  United  States  Biological  Survey  see 
McAtee  (1913). 

ALLEN,  C.  N. 

1881.     Songs  of  the  western  meadowlark.     Nutt.  Orn.  Club.  Bull.,  6, 

145-150. 
ALLEN,  J.  A. 

1876.     Progress  in  ornithology  in  the  United  States   during  the  last 

century.     Amer.  Nat.,  10,  536-550. 
1881.     Insectivorous  birds  in  their  relation  to  man.     Nutt.  Orn.  Club. 

Bull.,  6,  22-27. 

1912.     Remarks  on  the  case  of  Roosevelt  vs.  Thayer,  with  a  few  inde- 
pendent  suggestions   on  the   concealing   coloration   question. 
Auk,  29,  489-507. 
ANONYMOUS. 

1909.  The  true  function  of  birds.     Sci.  Amer.  S.,  68,  266-267. 

1910.  The  control  of  the  Argentine  ant.     Univ.  of  Calif.  Publ.,  Agric. 

Exp.  Sta.  Bull.,  207,  51-82,  28  figs,  in  text. 

1911.  Consider  the  birds  of  the  air.     Craftsman,  21,  372-421. 
ARCHIBALD,  C.  F. 

1907.     Wild  birds,  useful  and  injurious.     Jour.  Eoyal  Agric.  Soc.  Eng., 
68,  17-32,  7  figs,  in  text. 

AUDUBON,  J.   J. 

1831-39.     Ornithological    biography    (A.    Black,    Edinburgh),    5    vols., 

many  woodcuts  in  text. 
AUGHEY,  S. 

1878.     Notes  on  the  nature  of  the  food  of  birds  of  Nebraska.     U.  S. 

Entom.  Comm.,  First  Rep.,  Append.  2,  pp.  13-62. 
BAER,  W. 

1909.  Untersuchungsergebnisse   von   Mageninhalt    sachsischen   Vogel. 

Ornith.  Monatschr.  Magdeburg,  34,  33-34. 
BASKETT,  J.  N. 

1910.  Birds  and  insect  pests.     Sci.  Amer.,  103,  379. 
BARROWS,  W.  B. 

1912.  Michigan    bird    life    (Mich.    Agric.    College,    Lansing,    Mich.), 

xiv  +  822,  70  pis.,  151  figs,  in  text. 
BEAL,  F.  E.  L. 

1894.     The  meadowlark  and  Baltimore  oriole.    U.  S.  Dept.  Agric.  Year- 
book, 1894,  419-430,  2  figs,  in  text. 

1897.     Eevent  investigations  of  the  food  of  European  birds.     Auk,  14, 
8-14. 

1906.  Birds  as  conservators  of  the  forest.     State  of  N.   Y.   Forest, 

Fish,  and  Game  Comm.  Eep.,  1902-1903,  236-274,  14  pis. 

1907.  Birds  of  California  in  relation  to  the  fruit  industry.     Part  I, 

U.  S.  Dept.  Agric.,  Div.  Biol.  Surv.  Bull.,  30,  1-100,  5  pis. 

1908.  The  relations  between  birds  and  insects.     U.   S.   Dept.   Agric. 

Yearbook,  1908,  343-350. 


1914]   Bryant:  Economic  Status  of  the  Western  Meadowlark     495 

1910.  Birds  of  California  in  relation  to  the  fruit  industry.     Part  II, 

U.  S.  Dept.  Agric.,  Div.  Biol.  Surv.  Bull.,  34,  1-96,  6  pis. 
1912.     Food  of  our  more  important  flycatchers.     U.  S.  Dept.  Agric.  Bur. 

Biol.  Surv.  Bull.,  44,  1-67,  5  pis. 
BEEBE,  C.  W. 

1906a.  The   bird    (Henry   Holt   &   Co.,   N.   Y.),   xi  +  496,   371    figs,   in 

text.    The  food  of  birds,  pp.  142-164. 
1906b.  Oddities  of  bird  diet.     Outing,  47,  227-228. 
BENDIRE,  C.  E. 

1892.     Life-histories  of  North  American  birds.     Smithson.  Inst.  Con- 

trib.  Knowl.,  1,  ix  -f  445,  12  pis. 
1895.     Life-histories  of  North  American  birds.     Ibid.,  2,  ix  +  518,   7 

pis. 
BLANCHAN,  N. 

1903.     How   to    attract    the   birds    (Doubleday,   Page    &   Co.,   N.    Y.), 

1-224,  many  figs,  in  text. 
BOWDISH,  B.  S. 

1906.     The  rapid  growth  of  birds.     Sci.  Amer.  S.,  61,  25373-25374. 
BRYANT,  H.  C. 

1911.  The  relation  of  birds  to  an  insect  outbreak  in  northern  Cali- 

fornia during  the  spring  and  summer  of  1911.     Condor,  13, 

195-208,  4  figs,  in  text. 
1912a.  The  economic  status  of  the  meadowlark  in  California.     Mon. 

Bull.  State  Comm.  Hort.   (Sacramento,  Calif.),  1,  226-231,  2 

figs,  in  text,  2  tables. 
1912b.  The    Lewis    woodpecker — A    destroyer    of    almonds.      Ibid.,    1, 

362-366,  2  figs,  in  text. 
1912c.  Native  birds  and  mammals.    Western  Field  (Los  Angeles,  Calif.), 

19,  432-435,  2  figs,  in  text. 
1912d.  Birds  in  relation  to  a  grasshopper  outbreak  in  California.    Univ. 

Calif.  Publ.  Zool.,  11,  1-20. 
1912e.  Some  insects  and  other  arthropods  in  the  diet  of  the  western 

meadowlark.    Pomona  College  Jour.  Entomology  (Claremont, 

Calif.),  4,  807-809. 
1912f.  The    numbers    of    insects    destroyed    by    western    meadowlarks 

(Sturnella  neglecta).     Science,  36,  873-875. 
1913a.  Why  non-game  birds  should  be  protected.     Western  Wild  Life 

Call  (Pub.  by  Calif.  Associated  Societies  for  the  Conservation 

of  Wild  Life,  Berkeley,  Calif.),  no.  1,  pp.  8-11,  4  figs,  in  text. 
1913b.  The  economic  value  of  the  western  meadowlark  in  California. 

Univ.  Calif.  Publ.,  Agric.  Exp.  Sta.  Bull.,  236,  1-16,  7  figs. 

in  text. 

1913c.  Investigation  of  the  economic  status  of  non-game  birds.     Re- 
print from  Game  Bull.   no.   1,   Calif.   State  Fish   and  Game 

Comm.    (State  Printing  Office,   Sacramento,   Calif.),   1-19,  8 

figs,  in  text. 
1913d.  The  results  of  some  miscellaneous  stomach  examinations.     The 

Condor,  15,  92. 
BRYANT,  W.  E. 

1893.     Notes  on  tne  food  of  birds.    Zoe,  4,  54-58. 


496  University  of  California  Publications  in  Zoology    [VOL. 11 

CHAPMAN,  F.  M. 

1903.  The  economic  value  of  birds  to  the  state.     State  of  New  York 

Forest,  Fish,  and  Game  Comm.,  1903,  1-66,  12  pis. 

1907.  Bird  life    (Appleton   &   Co.,   N.   Y.),  xii  -f  88,   75   pis.,   25   figs. 

in  text. 

1908.  Camps  and  cruises  of  an  ornithologist  (Appleton  &  Co.,  N.  Y.), 

xvi  -f  432,  illus. 

1909.  A  study  of  the  genus  Sturnella.     Amer.  Mus.  Nat.  Hist.  Bull., 

13,  297-320,  8  figs,  in  text. 
CHEENELHAZA,  S.  C.  VON 

1909.  Beitrage    zur    Nahrungsfrage    unserer    carnivoren    Vogelwelt. 

Aquila,  Budapest,  16,  145-155. 
CLELAND,  J.  B. 

1910.  Examination  of  contents  of  stomachs  and  crops  of  some  Austra- 

lian birds.     Emu,  9,  219-226.     Agric.  Gaz.  N.  S.  Wales,  21, 
73-156. 

COLLINGE,  W.  E. 

1912.     The  food  of  nestling  birds.     Jour.  Bd.  Agric.,  London,  19,  460- 

465. 
COUES,  E. 

1874.     Birds    of    the    northwest    (Gov.    Printing    Office,    Washington), 

U.  S.  Geol.  Surv.  of  Terr.,  Misc.  Publ.,  3,  xi  +  791. 
COWARD,  T.  A. 

1912.     The  migration  of  birds  (Univ.  Press,  Cambridge;  G.  P.  Putnams 

Sons,  N.  Y.),  1-137,  4  maps. 
COWAN,  J.  C. 

1911.  Eefuges  for  birds.    Overland,  57,  637-642. 
GEARY,  L.  E. 

1911.     A  text-book  of  field  zoology   (Blakiston's  Sons  &  Co.,  Phila., 

Pa.),  1-364,  117  figs,  in  text. 
CSIKI,  E. 

1909.     Positive  Daten  iiber  die  Nahrung  unserer  Vogel.    Aquila,  Buda- 
pest, 16,  139-144. 
DAEHNE 

1909.     Schmetterlingsfeinde  aus  der  Klasse  der  Vogel.     Zs.  Naturw., 

Stuttgart,  81,  184-187. 
DAWSON,  W.  LEON,  and  BOWLES,  J.  H. 

1909.     The  birds  of  Washington  (Occidental  Publ.  Co.,  Seattle,  Wash., 

2  vols.,  xv  +  997,  44  pis.,  300  figs,  in  text. 
DEWAR,  D.,  and  FINN,  F. 

1909.     The    making   of    species    (J.   Lane    &    Co.,    N.    E.),    xix  -f-  400, 
10  pis. 

DUGMOEE,  A.  R. 

1907.     Birds  as  insect  destroyers.     Country  Life,  11,  384-390. 
BUTCHER,  WM. 

1904.  Report  of  the  A.  O.  U.  committee  on  the  protection  of  North 

American  birds  and  of  the  national  committee  of  Audubon 
societies.     Auk,  21,  Supp.,  97-208. 


1914]   Bryant:  Economic  Status  of  the  Western  Meadowlark     497 

ELLIOT,  D.  C. 

1864.     The  game  birds  of  the  United  States.    U.  S.  Dept.  Agric.  Eep., 

1864,  356-385,  5  pis. 
FINLEY,  W.  L. 

1907.     American  birds    (Chas.   Scribner's  Sons,  N.  Y.),  xvi  +  256,  127 
figs,  in  text. 

1909.  Study  of  birds  and  their  economic  value:  Some  common  birds 

of  Oregon.    Fourth  Biennial  Keport  01  the  State  Biologist  of 

Oregon,  Append.,  pp.  3-24. 
FISHER,  A.  K. 

1887.     Food  of  hawks  and  owls.    U.  S.  Dept.  Agric.,  Ann.  Report,  1887, 

402-422. 
1893.     The  hawks  and  owls  of  the  United  States  in  their  relation  to 

agriculture.     U.   S.   Dept.   Agric.,  Div.   Biol.   Surv.  Bull.,   3, 

1-210,  26  pis. 
FLAGG,  W. 

1862.     Utility  of  birds.    Mass.  Agric.  Eep.,  1861,  70-78. 
FLORENCE,  L. 

1912.     The  food  of  birds.     Trans.  Highland,  and  Agric.  Soc.  Scotland, 

24,  180-219. 
FOLSOM,  J.  W. 

19C9.     Entomology  with  especial  reference  to  its  biological  and  eco- 
nomic aspects   (P.  Blakiston's  Sons  &  Co.,  Phila.,  Pa.),  vii 

+  485,  5  pis.,  300  figs,  in  text. 
FORBES,  S.  A. 

1880.     On  some  interactions  of  organisms.     Bull.  111.  State  Lab.  Nat. 

Hist.,  1,  3-17. 
1882.     The  ornithological  balance-wheel.     Trans.  111.  State  Hort.  Soc., 

new  ser.,  15,  120-131. 
1885.     The  regulative  action  of  birds  upon  insect  oscillations.   111.  State 

Lab.  Nat.  Hist.  Bull.,  1,  3-32. 
1903.     The  food  of  birds.    Ibid.,  1,  86-161. 

1907.  An   ornithological   cross-section   of  Illinois.     Ibid.,  7,  305-335, 

12  tables. 

1908.  The  midsummer  bird  life  of  Illinois:  A  statistical  study.    Amer. 

Nat.,  42,  505-519. 
FORBUSH,  E.  H. 

1908a.  Birds  as  protectors  of  orchards.     Mass.  State  Bd.  Agric.  Bull., 

July,  1895,  1-19. 
1908b.  Two  years  with  the  birds  on  a  farm.     Ibid.,  June,  1908,  1-44, 

8  figs. 
1908c.  Useful  birds  and   their  protection    (3rd   ed.,   Mass.   State   Bd. 

of  Agric.,  Boston),  xx  +  437,  56  pis.,  171  figs,  in  text. 

1910.  Third  annual  report  of  the  State  Ornithologist.     Mass.   State 

Bd.  of  Agric.  Eep.,  58,  167-197. 
GAUT,  H.  L. 

1906.     What  do  birds  eat?    Sci.  Amer.,  95,  379. 
GENTRY,  T.  G. 

1876.     Life-histories  of  the  birds  of  Pennsylvania  (Pub.  by  the  author, 
Phila.),  vol.  2,  xiv  +  399. 


498          University  of  California  Publications  in  Zoology    [V°L. 11 

GILMOUR,  JOHN 

1896.     Bird  investigation:  An  inquiry  concerning  the  relation  of  cer- 
tain birds  to  the  agricultural  interest,  as  shown  by  their  diet. 
Jour.  Highland.  Agric.  Soc.,  1896,  21-112. 
GLADDEN,  GEO. 

1912.     Shall  Uncle  Sam  protect   the  birds.     Amer.   Eev.   of  Eev.,  46, 

705-715,  20  figs,  in  text. 
Go s SARD,  H.  A. 

1908.     Spring  practice   in   economic   zoology.     Ohio   Agric.   Exp.    Sta. 

Bull.,  198,  15-88,  Appendix  viii. 
GRESCHIK,  EUGEN 

1910.  Magen-  und  Gewolluntersuchungen  unserer  unheimischen  Baub- 

vogel.     I.  Mitteilung.     Aquila,  17,  1-13. 

1911.  II.  Eulen.     Ibid.,  18,  111-177,  6  figs,  in  text. 
GUNNING,  j.  W. 

1908.     Locusts  birds.     Transvaal  Agric.  Jour.,  6,  527-530. 
HAMMOND,  JOHN 

1912.  An  investigation  concerning  the  food  of  certain  birds.     Jour. 

of  Agric.  Soc.  London,  4,  380-407. 
HENDERSON,  J. 

1913.  The  practical  value  of  birds.     Univ.  of  Colo.  Bull.,  13,  1-48. 
HEN  SHAW,  H.  W. 

1907.  Does  it  pay  the  farmer  to  protect  the  birds?    U.  S.  Dept.  Agric. 

Rep.,  1907,  165-178. 

1908.  The  policemen  of  the  air:  An  account  of  the  Biological  Survey 

of   the   Department    of   Agriculture.      Nat.    Geog.    Mag.,    19, 
79-118,  38  figs,  in  text. 
HEWITT,  C.  G. 

1909.  An  inquiry  into  the  feeding  habits  of  British  birds.     British 

Ass.  Adv.  Sci.  Eep.,  1908,  734-735. 

HOLLRUNG,  M. 

1906.     Beitrage    zur    Bewertung    der    Saatkrahe    auf    Grund    von    11- 
jahrigen  Magenuntersuchungen.     Landwirtschaft.  Jahrb.  35, 
579-620,  1  fig.  in  text. 
HOLMES,  E. 

1857.     Birds  injurious  to  agriculture.     Eep.  U.  S.  Patent  Office,  Agric., 

1856,  1857,  110-160,  pis.  15-46. 
HOOPER,  C.  H. 

]908.     The    commoner   birds   of   our   gardens,   their   habits   and    food. 

Jour.  Eoyal  Hort.  Soc.  London,  33,  427-450,  1  fig.  in  text. 
HORN  AD  AY,  W.  T. 

1913.     Our    vanishing    wild    life     (Chas.     Scribner's     Sons,     N.     Y.), 
xv  +  411,    many   illus.      The    economic    value    of    birds,    pp. 
211-233. 
JENKS,  J.  W.  P. 

1860.     The   food   of   the   robin.      Jour.    Proc.   Mass.    Hort.    Soc.,   1859, 
151-154. 


1914]   Bryant:  Economic  Status  of  the  Western  Meadowlark     499 

JUDD,  S.  D. 

1897.  Methods  in  economic  ornithology,  with  especial  reference  to  the 
catbird.  Amer.  Nat.,  31,  392-397. 

1899.  The  efficiency  of  some  protective  adaptations  in  securing  insects 

from  birds.     Ibid.,  33,  461-484. 

1900.  Food  of  nestling  birds.    U.  S.  Dept.  Agric.  Yearbook,  1900,  411- 

436. 

1901.  The  relation  of  sparrows  to   agriculture.     U.   S.   Dept.   Agric., 

Div.  Biol.  Surv.  Bull.,  15,  1-98,  4  pis.,  19  figs,  in  text. 

1902.  Birds  of  a  Maryland  farm.     Ibid.,  17,  1-116,  16  pis.,  41  figs,  in 

text. 
KALBFUS,  JOSEPH 

1911.     Wild  bird  protection.     Penn.  State  Bd.  of  Game  Comm.  Bull., 

2,  1-23,  12  figs,  in  text. 
KELLOGG,  V.  L. 

1908.     American  insects   (Holt  &  Co.,  N.  Y.),  xiv  -f  694,  812  figs,  in 

text. 
KING,  F.  H. 

1883.     Economic   relations   of  Wisconsin   birds.      Geology   of   Wis.,   1, 

441-610,  41  figs,  in  text. 
KIRBY,  WM.,  and  SPENCE,  WM. 

1858.     An    introduction    to    entomology    (7th    ed.,    Longman,    Brown, 
Green,  Longmans,  and  Eoberts,  London),  xxviii  -f-  607.    Inter- 
action of  organisms,  p.  145. 
KNIGHT,  W.  C. 

1902.     The  birds  of  Wyoming.    Wyo.  Agric.  Exp.  Sta.  Bull.,  55,  1-174. 
LEISEWITZ,  W. 

1906.  Untersuchungen  iiber  den  Nahrung  einiger  land-  und  forstwirt- 

schafthch  wichtigen  Vogelarten.     Verhand.  der  Orn.  Gesell- 
schaft  in  Bayern,  6,  194-203. 

1907.  Ueber  neuere  Untersuchungen  zur  Kenntnis  der  Nahrung  der 

Vogel.    Ibid.,  7,  265-274. 

1911.  Untersuehungen   des   Inhalts   von   Kaubvb'gelmagen.     Ibid.,   10, 

156-182. 
LE  BARON,  W. 

1855.     Observations  upon  some  of  the  birds  of  Illinois  most  interesting 
to  the  agriculturist.     Trans.  111.  State  Agric.  Soc.,  1853-1854, 
559-565. 
LOCY,  W.  A. 

1910.     Biology  and  its  makers  (Henry  Holt  &  Co.,  N.  Y.),  xxvi  +  469, 

123  figs,  in  text. 
MASON,  C.  W.,  and  LEFROY,  H.  M. 

1912.  Preliminary  report  on  food  of  Indian  birds.     Mem.  Dept.  Agric. 

India,  Entom.  Ser.,  3,  1-371. 

MCATEE,  W.  L. 

1906.     A  buried  treasure  of  economic  ornithology.     Sci.,  24,  308-312. 

1912a.  The  experimental  method  of  testing  the  efficiency  of  warning 
and  cryptic  coloration  in  protecting  animals  from  their  ene- 
mies. Proc.  Acad.  Nat.  Sci.  Phila.,  1912,  281-364. 


500  University  of  California  Publications  in  Zoology    [VOL. 11 

1912b.  Methods  of  estimating  the  contents  of  bird  stomachs.  Auk,  29, 
449-464. 

1912c.  Eeview  of  Bryant's  "The  relation  of  birds  to  an  insect  out- 
break in  northern  California  during  the  spring  and  summer 
of  1911.  Condor,  14,  45-46. 

1913.     Index  to  papers  relating  to  the  food  of  birds.    U.  S.  Dept.  Agric., 

Bur.  Biol.  Surv.  Bull.,  43,  1-69. 
MERRIAM,  T.  A. 

1896.  How  birds  effect  the  farm  and  garden.     Forest  and  Stream,  47, 

103,  123,  141. 
.MINOT,  CHAS.  A. 

1911.     The  method  of  science.     Sci.,  33,  119-131. 
NASH,  CHAS.  W. 

1909.     The  birds  of  Ontario  in  relation  to  agriculture.     (4th  ed.),  Ont. 

Dept.  Agric.  Bull.,  173,  1-75,  pis. 
PEABODY,  P.  B. 

1897.  Song   notes    and    nesting    habits    of   the    western    meadowlark. 

Osprey,  1,  139-141,  1  fig.  in  text. 
PALMER,  T.  S. 

1899.     A  review  of  economic  ornithology.     U.  S.   Dept.  Agric.   Year- 
book, 1899,  259-292. 
PEARSON,  T.  G. 

1909.     Some  common  birds  on  the  farm.     N.  C.  Dept.  Agric.,  Special 

Bull.,  Supp.  Sept.  Bull.,  1909,  1-28,  5  figs,  in  text. 
1913.     Economic  value  of  birds.     Craftsman,  23,  432-438. 
PENNOCK,  CHAS.  J. 

1904.     Some  of  our  useful  birds.     Dela.  State  Bd.  of  Agric.  Bull.,  5, 

1-16,  6  figs,  in  text. 
PERRIS,  E. 

1880.     Birds  vs.  insects.     (Translated  by  S.  A.  Forbes.)     Amer.  Entom., 

1,  69-72,  96-100. 
POCOCK,  E.  I. 

1893.     Further  notes  and  observations  on  the  instincts  of  some  com- 
mon English  spiders.     Nature,  49,  61-63. 
POULTON,  E.  B. 

1887.     The  experimental  proof  of  the  protective   value   of  color  and 
color  markings  in   insects   in   reference   to   their   vertebrate 
enemies.    Proc.  Zool.  Soc.  London,  1887,  191-274,  5  tables. 
1890.     The   colors   of   animals    (Kegan   Paul,   Trench,   Triibner   &   Co., 
London),  xiii  +  360. 

1898.  Natural  selection,  the  cause  of  mimetic  resemblance  and  com- 

mon warning  colors.     Jour.  Linn.  Soc.  London,  Zool.  Ser.,  26, 
558-612,  pis.  40-44. 

1908.     Essays   on   evolution    (Clarendon   Press,   Oxford),   xlviii  -+-  479. 
PRAEGER,  WM.  E. 

1899.  Birds  in  horticulture.     Trans.  111.  State  Hort.  Soc.,  New  Ser., 

33,  1-11. 


1914]   Bryant:  Economic  Status  of  the  Western  Meadowlark     501 

PREVOST,  M.  F. 

1858.     A  memoir  on  the  alimentary  regimen  of  birds  presented  to  the 
Imperial  Zoological  Society  of  Paris.     (Translated  by  J.  W. 
P.  Jenks.)     Trans.  Mass.  Hort.  Soc.,  1859,  109-114. 
PYCBAFT,  W.  P. 

1910.     A    history   of   birds    (Methuen      Co.,   London),    xxxi  -f  458,    36 

pis.,  50  figs,  in  text. 
KOGERS,  J.  E. 

1910.  Birds  that  save  our  crops.     Country  .Life,  18,  250-254. 
ROOSEVELT,  T. 

1911.  Revealing    and    concealing    coloration   in    birds   and    mammals. 

Amer.  Mus.  Nat.  Hist.  Bull.,  30,  119-231. 
RORIG,  G. 

1903a.  Studien  iiber  die  wirtschaftlichen  Bedeutung  der  insekton-fres- 

senden  Vogel.     Berlin  Arb.  biol.  Anst.,  4,  1-50. 
1903b.  Untersuchungen  iiber  die  Nahrung  unserer  heimischen  Vogel  mit 

besonderer  Beriicksichtigung  der  Tag-  und  Naehtraubvb'gel. 

Ibid.,  51-120,  3  Tafeln. 

1909.  Magen-  und  Gewolluntersuchungen  heimischen  Raubvogel.     Ber- 

lin biol.  Anst.,  7,  473-520. 
SAMPLE,  O.  H. 

1910.  The  conservation  of  birds.     Outlook,  95,  669-676,  illus. 

SCLATER,  W.  L. 

1912.  A  history  of  the  birds  of  Colorado   (Witherby  &  Co.,  London), 

xxiv  -f-  576,  16  pis.,  1  map. 
SHUFELDT,  R.  W. 

1887.     On  the  skeleton  in  the  genus  Sturnella,  with  osteological  notes 
upon    other    North    American    Icteridae    and    the    Corvidae. 
Jour,  of  Anat.  and  Physiol.,  22,  309-350,  2  pis. 
SURFACE,  H.  A. 

1908.     Birds  and  horticulturists.    Penn.  State  Dept.  Agric.,  Mon.  Bull. 

Div.  of  Zool.,  5,  299-320,  3  pis.,  7  figs,  in  text. 
1905.     The  economic  value  of  our  native  birds.     Ibid.,  3,  240-267,  6 

pis. 
SULLIVAN,  R.  H. 

1910.     The  economic  value  of  bird  life.     Agric.  Educ.,  3,  1-47,  30  figs, 
in  text. 

SWYNNERTON,  C.  F.  M. 

1912.     Remarks  on  the  stomach  contents  of  birds.     Ibis,  6,  635-640. 
TITCHENER,  E.  B.,  and  FINN,  F. 

1889.     Comparative  palatability  of  insects,  etc.     Nature,  42,  571-572. 

1891.  Ibid.,  44,  540. 

1892.  Ibid.,  45,  53. 
THEOBALD,  F.  V. 

1908.     Economic  ornithology   in  relation   to   agriculture,   horticulture, 

and  forestry.     Sci.  Progress,  2,  263-283. 
TREADWELL,  D. 

1859.     The  food  of  young  robins.    Proc.  Boston  Soc.  Nat.  Hist.,  6,  396- 
399.    Review  U.  S.  Patent  Office  Rep.  on  Agric.,  1860,  88-89. 


502  University  of  California  Publications  in  Zoology    [VOL.  11 

WARREN,  B.  H. 

1888.     Eeport   of  the  birds  of  Pennsylvania    (E.  K.   Meyers,   Harris- 
burg,  Pa.),  xii  +  260,  50  pis. 
WEED,  C.  M. 

1902.  A    partial    bibliography    of    the    economic    relations    of    North 

American  birds.     N.  H.  College  Agric.  Exp.  Sta.  Tech.  Bull., 
5,  139-179. 
1910.     Farm    friends    and    farm    foes    (D.    C.    Heath    &    Co.,    Boston, 

Mass.),  xi  +  334,  1  map,  160  figs,  in  text. 
WEED,  C.  M.,  and  DEARBORN,  N. 

1903.  Birds  in   their  relation   to  man    (Lippincott,   Phila.,  Pa.),   viii 

+  380,  illus. 
WHEELOCK,  I.  G. 

1905.  Eegurgitative  feeding  of  nestlings.     Auk,  22,  54-71. 

1906.  Nesting  habits  of  the  green  heron.    Ibid.,  23,  432-436. 
WIER,  J.  J. 

1869.     On  some  insects  and  insectivorous  birds,  and  especially  on  the 
relation  between  color  and  the   edibility  of  Lepidoptera   and 
their  larvae.     Trans.  Ent.  Soc.  London,  1869,  21-26. 
WILCOX,  E.  V. 

1892.     The  food  of  the  robin.     Ohio  Agric.  Exp.   Sta.  Bull.,  43,   115- 

131. 
WILSON,  A. 

1808-14.     American   Ornithology    (Bradford    &   Inskeep,    Phila.,    Pa.), 
9  vols.,  many  pis.  and  figs. 

WOODWORTH,  C.  W. 

1902.  Grasshoppers  in  California.     Univ.  of  Calif.  Publ.,  Agric.  Exp. 

Sta.  Bull.,  142,  1-36,  17  figs,  in  text. 

1903.  A  list  ot  the  insects  of  California.     (Pub.  by  the  author,  Berke- 

ley, Calif.),  pp.  1-80. 


EXPLANATION  OF  PLATES 

PLATE  21 

Fig.  1. — Holes  bored  by  western  meadowlarks  in  obtaining  kernels  of 
grain  in  sprouting  grain  field.  Photograph  by  H.  C.  Bryant  taken  at 
Lathrop,  San  Joaquin  County,  California,  February  28,  1912. 

Fig.  2. — Sprouted  grain  pulled  up  by  western  meadowlarks  in  grain 
fields  at  Acampo  and  Lathrop,  San  Joaquin  County,  California.  The 
kernels  have  been  crushed  in  the  bill  to  obtain  the  "milk". 


[504] 


UNIV.   CALIF.    PUBL.   ZOOL.  VOL, 


[BRYANT]    PLATE  21 


Fig.  1 


Fig.  2 


PLATE  22 

Fig.  3. — Photograph  of  the  stomach  contents  of  a  western  meadowlark 
taken  in  a  grain  field  at  El  Toro,  Orange  County,  Calirornia,  May  5,  1911. 
The  stomach  contained  19  oat  kernels,  oat  hulls,  and  parts  of  2  small 
ground  beetles. 

Fig.  4. — Photograph  of  twelve  pairs  of  mandibles  of  the  common  cricket 
(Gryllus  pennsylvanidus)  taken  from  the  stomach  of  a  western  meadowlark 
collected  in  a  grain  field  at  El  Toro,  Orange  County,  California,  April  18, 
1911. 


[506] 


UNIV.   CALIF.    PUBL.   ZOOL.   VOL.    II 


[BRYANT]    PLATE  22 


Fig.  3 


t 


I 


Fig.  4 


PLATE  23 

Fig.  5. — Photograph  of  the  stomach  contents  of  a  western  meadowlark 
taken  at  Eed  Bluff,  Tehama  County,  California,  February  4,  1911.  Stom- 
ach contained  66  cutworms,  18  small  ground  beetles,  2  beetle  larvae,  2 
small  spiders,  and  18  weed  seeds. 

Fig.  6. — Photograph  of  western  meadowlark  killed  while  carrying  cut- 
worms in  its  bill. 


[508] 


UNIV.   CALIF.    PUBL.  ZOOL.   VOL.    II 


[BRYANT]    PLATE  23 


-  i€ 

-•*  < 

-  ^  AI 

-  '  i  --   > 

:1  v"?   i*/C 


Fig.  6 


PLATE  24 

Fig.  7. — Photograph  of  stomach  contents  of  a  western  meadowlark 
taken  in  a  barley  field  at  El  Toro,  Orange  County,  California,  May  3, 
1911.  The  stomach  contained  2  cutworms,  44  ground  beetles  (Calathus 
ruficollis'),  2  flies,  1  spider,  and  13  fly  pupae  (Syrphus  sp.). 

Fig.  8. — Photograph  of  the  stomach  contents  of  a  western  meadowlark 
collected  at  Big  Pine,  Inyo  County,  California,  April  19,  1911.  The  stom- 
ach contained  13  cutworms,  26  elaterid  beetles  (Drasterius  sp.),  and  10 
small  ground  beetles  (Amara  sp.). 


[510] 


UNIV.   CALIF,    PUBL   ZOOL.    VOL, 


[BRYANT]    PLATE  24 


»     r  •  t  4  I  t 


I 


Fig.  7 


Fig. 


UNIVERSITY  OF  CALIFORNIA  PUBLICATIONS— (Continued) 

rol.  7.     (Contributions  from  the  Museum  of  Vertebrate  Zoology.) 

1.  Two  New  Owls  from  Arizona,  with  Description  of  the  Juvenal  Plum- 

age of  Strix  occidentalis  occidentalis  (Xantus),  by  Harry  S.  Swarth. 

Pp.  1-8.     May,  1910      „ „ 10 

2.  Birds  and  Mammals  of  the  1909  Alexander  Alaska  Expedition,  by 

Harry  S.  Swarth.  Pp.  9-172;  plates  1-6;  3  text-figures.  January,  1911.    1.50 

3.  An  Apparent  Hybrid  in  the  Genus  Dendroica,  by  Walter  P.  Taylor. 

Pp.  173-177.     February,  1911 05 

4.  The  Linnet  of  the  Hawaiian  Islands:   a  Problem  in  Speciation,  by 

Joseph  Grinnell.    Pp.  179-195.    February,  1911  15 

5.  The  Modesto  Song  Sparrow,  by  Joseph  Grinnell.    Pp.  197-199.    Feb- 

ruary, 1911  '. 05 

6.  Two  New  Species  of  Marmots  from  Northwestern  America,  by  H.  S. 

Swarth.    Pp.  201-204.    February,  1911  05 

7.  Mammals  of  the  Alexander  Nevada  Expedition  of  1909,  by  Walter  P. 

Taylor.    Pp.  205-307.    June,  1911  • 1.00 

8.  Description  of  a  New  Spotted  Towhee  from  the  Great  Basin,  by  J. 

Grinnell.    Pp.  309-311.    August,  1911  05 

9.  Description  of  a  New  Hairy  Woodpecker  from  Southeastern  Alaska,  by 

H.  S.  Swarth.    Pp.  313-318.    October,  1911  . 05 

10.  Field  Notes  on  Amphibians,  Reptiles  and  Birds  of  Northern  Humboldt 
County,  Nevada,  with  a  Discussion  of  Some  of  the  Faunal  Features 
of  the  Region,  by  Walter  P.  Taylor.  Pp.  319-436,  plates  7-12. 

February,  1912 1.00 

Index,  pp.  437-446. 
Vol.  8.      1.  The  Vertical  Distribution  of  Eucalanus  elongatus  in  the  San  Diego 

Region  during  1909,  by  Calvin  O.  Esterly.    Pp.  1-7.    May,  1911 10 

2.  New  and  Rare  Fishes  from  Southern  California,  by  Edwin  Chapin 

Starks  and  William  M.  Mann.    Pp.  9-19,  2  text-figures.    July,  1911.      .10 

3.  Classification  and  Vertical  Distribution  of  the  Chaetognatha  of  the  San 

Diego  Region,  Including  Redescriptions  of  Some  Doubtful  Species  of 

the  Group,  by  Ellis  L.  Michael.  Pp.  21-186,  pis.  1-8.  December,  1911.    1.75 

4.  Dinoflagellata  of  the  San  Diego  Region,  IV.  The  Genus  Gonyaulax,  with 

Notes  on  Its  Skeletal  Morphology  and  a  Discussion  of  Its  Generic 
and  Specific  Characters,  by  Charles  Atwood  Kofoid.  Pp.  187-286, 
plates  9-17. 

5.  On  the  Skeletal  Morphology  of  Gonyaulax  catenata   (Levander),  by 

Charles  Atwood  Kofoid.    Pp.  287-294,  plate  18. 

6.  Dinoflagellata  of  the  San  Diego  Region,  V.  On  Spiraulax,  a  New  Genus 

of  the  Peridinida,  by  Charles  Atwood  Kofoid.    Pp.  295-300,  plate  19. 

Nos.  4,  5,  and  6  in  one  cover.    September,  1911  1.50 

7.  Notes  on  Some  Cephalopods  in  the  Collection  of  the  University  of  Cali- 

fornia, by  S.  S.  Berry.    Pp.  301-310,  plates  20-21.    September,  1911.      .10 

8.  On  a  Self-Closing  Plankton  Net  for  Horizontal  Towing,  by  Charles 

Atwood  Kofoid.    Pp.  311-348,  plates  22-25. 

9.  On  an  Improved  Form  of  Self-closing  Water-bucket  for  Plankton  In- 

vestigations, by  Charles  Atwood  Kofoid.    Pp.  349-352. 

Nos.  8  and  9  in  one  cover.    November,  1911 40 

Index,  pp.  353-357. 

Vol.  9.  1.  The  Horned  Lizards  of  California  and  Nevada  of  the  Genera  Phryno- 
soma  and  Anota,  by  Harold  C.  Bryant.  Pp.  1-84,  plates  1-9.  Decem- 
ber, 1911 70 

2.  On  a  Lymphoid  Structure  Lying  Over  the  Myelencephalon  of  Lepisos- 

teus,  by  Asa  C.  Chandler.    Pp.  85-104,  plates  10-12.    December,  1911.      .25 

3.  Studies  on  Early  Stages  of  Development  in  Rats  and  Mice,  No.  3,  by 

E.  L.  Mark  and  J.  A.  Long.  The  Living  Eggs  of  Rats  and  Mice  with 
a  Description  of  Apparatus  for  Obtaining  and  Observing  Them  (Pre- 
liminary paper),  by  J.  A.  Long.  Pp.  105-136,  plates  13-17.  February, 
1912 , SO 

4.  The  Marine  Biological  Station  of  San  Diego,  Its  History,  Present  Con- 

ditions, Achievements,  and  Aims,  by  Wm.  E.  Ritter,  Pp.  137-248, 
plates  18-24,  and  2  maps.  March,  1912  _ 1.00 

5.  Oxygen  and  Polarity  in  Tubularia,  by  Harry  Beal  Torrey.    Pp.  249- 

251.     May,  1912  05 

6.  The  Occurrence  and  Vertical  Distribution  of  the  Copepoda  of  the  San 

Diego  Region,  with  particular  reference  to  Nineteen  Species,  by  Cal- 
vin O.  Esterly.  Pp.  253-340,  7  text-figures.  July,  1912  1.00 

7.  Observations  on  the  Suckling  Period  in  the  Guinea-Fig,  by  J.  Marion 

Read.    Pp.  341-351.    September,  1912  10 

8.  HaeckePs  Sethocephaliis  encecryphalus  (Radiolaria),  a  Marine  Ciliate, 

by  Charles  Atwood  Kofoid.    Pp.  353-357.    September,  1912  .05 

Index,  pp.  359-365, 


u.u.   ui_i  U\I_I_L  i 


NON-CIRCULATING  BOOK 


UNIVERSITY  OF  CALIFORNIA  LIBRARY 


